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JeL GENEVE JUIN 2004 ISSN 0035 - 418 X GO fanH 



REVUE SUISSE DE ZOOLOGIE 



TOME 111— FASCICULE 2 

Publication subventionnée par: 

Académie suisse des Sciences naturelles ASSN 

Ville de Genève 

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CHARLES LIENHARD 

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Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du 
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comparée, physiologie. 

Administration 

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REVUE SUISSE DE ZOOLOGIE 



TOME 111 -FASCICULE 2 

Publication subventionnée par: 

Académie suisse des Sciences naturelles ASSN 

Ville de Genève 

Société suisse de Zoologie 



VOLKER MAHNERT 
Directeur du Muséum d'histoire naturelle de Genève 

CHARLES LIENHARD 
Chargé de recherche au Muséum d'histoire naturelle de Genève 



Comité de lecture 

Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du 
Muséum de Genève et de représentants des instituts de zoologie des universités suis- 
ses. 

Les manuscrits sont soumis à des experts d'institutions suisses ou étrangères selon le 
sujet étudié. 

La préférence sera donnée aux travaux concernant les domaines suivants: biogéo- 
graphie, systématique, évolution, écologie, éthologie, morphologie et anatomie 
comparée, physiologie. 

Administration 

MUSÉUM D'HISTOIRE NATURELLE 
1211 GENÈVE 6 

Internet: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm 



Prix de l'abonnement: 

SUISSE Fr. 225. - UNION POSTALE Fr. 230. - 

(en francs suisses) 

Les demandes d'abonnement doivent être adressées 

à la rédaction de la Revue suisse de Zoologie, 

Muséum d'histoire naturelle, C.P 6434, CH-1211 Genève 6, Suisse 



Revue suisse de Zoologie 111 (2): 239-256; juin 2004 



Systematics and phylogenetic relationships of Whip snakes 
{Hierophis Fitzinger) and Zamenis andreana Werner, 1917 
(Reptilia: Squamata: Colubrinae) 

Beat SCHÄTTI 1 & Paul MONSCH 2 

1 Apartado postal 383, San Miguel de Allende, Gto. 37700, Repüblica Mexicana. 

2 Teilstrasse 4a, CH-8400 Winterthur, Switzerland. 



Systematics and phylogenetic relationships of Whip snakes {Hierophis 
Fitzinger) and Zamenis andreana Werner, 1917. - Morphological and 
molecular data of Coluber (sensu lato) andreanus and all recognised species 
of the Palaearctic whip snake genus Hierophis Fitzinger are presented. 
Morphologically, Andreas' racer shows remarkable similarities to dwarf 
snakes {Eirenis spp., Pseudocyclophis persicus). Derived conditions of head 
and body pholidosis including dorsal scale reduction pattern and the number 
of apical pits separate Eirenis Jan and C. (s.l.) andreanus from Hierophis 
spp. Character states strongly support sister group relationship of dwarf 
snakes to Andreas' racer. Molecular analyses confirm monophyly of Hiero- 
phis spp., C. (s.l.) andreanus, and dwarf snakes of the genus Eirenis, and 
paraphyly of Hierophis auct. MtDNA sequences indicate a basal position of 
the eastern Palaearctic H. spinalis vis-à-vis the western species group (H. 
cypriensis, H. gemonensis, H. viridiflavus). These taxa belong to an early 
radiation within whip and dwarf snakes. The eastern Mediterranean H. 
caspius, H. jugularis, and H. schmidti appear to represent a paraphyletic 
grouping including C. (s.l.) andreanus (12S rDNA sequence data). 
Conflicting molecular and morphological results are discussed and the 
character phylogeny of external morphological features is re-assessed. 

Key-words: Coluber (s.l.) andreanus - Eirenis - Hierophis - morphology - 
osteology - hemipenis - transformation series - phylogeny - mtDNA. 

INTRODUCTION 

Without taking account of Oriental taxa (Utiger, 2002), Old World racers and 
their allies belong to at least eight different genera, i.e., Eirenis Jan, 1863, Hemerophis 
Schätti & Utiger, 2001, Hemorrhois Boie, 1826, Hierophis Fitzinger in Bonaparte, 
1834, Lytorhynchus Peters, 1862, Platyceps Blyth, 1860, Pseudocyclophis Boettger, 
1888, and Spalerosophis Jan, 1865 (Schätti 1986, 1987; Schätti & Utiger, 2001). The 
latter genus is the sister taxon of Hemorrhois Boie. Together with Platyceps Blyth, they 
make up a monophyletic group among Old World racers. 



Manuscript accepted 03.11.2003 



240 B. SCHÄTTI & P. MONSCH 

The Palaearctic whip snake (or racer) genus Hierophis Fitzinger belongs to a dis- 
tinct evolutionary lineage within Afrotropical, Palaearctic, and Saharo-Sindian racers. It 
is considered to include H. caspius (Gmelin, 1789), H. cypriensis (Schätti, 1985), H. 
gemonensis (Laurenti, 1768), H. jugularis (Linnaeus, 1758), H. schmidti (Nikolskij, 
1909), H. spinalis (Peters, 1866), and H. viridiflavus (Lacépède, 1789), the type species 
(Schätti, 1987, 1988, 1993; Schätti & Utiger, 2001; Utiger & Schätti, 2004). 

Zamenis andreana Werner, 1917 from the southern Zagros Mountains in Iran is 
a poorly known and enigmatic species characterised by a number of distinctive 
external morphological features typically encountered in dwarf snakes of the genus 
Eirenis Jan. Schätti (2001) thought that Andreas' racer is a representative of an early 
evolutionary lineage among Palaearctic racers. Schätti & Utiger (2001) placed the 
taxon in their incertae sedis section, i.e., Coluber sensu lato, stressing the necessity of 
comparison with Eirenis spp. 

A preliminary molecular analysis (Schätti & Utiger, 2001) generated low boot- 
strap values for the European {Hierophis gemonensis, H. viridiflavus) and eastern 
Mediterranean (H. caspius, H. jugularis, H. schmidti) whip snakes. On the basis of 
mtDNA sequence data, Eirenis modestus (Martin, 1838) turned out to be the sister 
taxon of the latter species group, thus rendering Hierophis auct. paraphyletic. The 
reality of two whip snake groups was also confirmed with cytochrome b sequences 
(Nagy et al., 2000). 

Using external morphological characters, vertebra (only Hierophis spp.) and 
hemipenis features, as well as molecular methods, this study investigates the phylo- 
genetic relationships of Zamenis andreana Werner, the systematic composition of 
Hierophis, and the morphological demarcation of whip snakes vis-à-vis dwarf snakes, 
i.e., Eirenis spp. and Pseudocyclophis persicus (Anderson, 1872). 

MATERIAL AND METHODS 

Methods and definitions of terms used in the text are explained in Schätti (1987, 
1988). For the purpose of this study, the number of anterior suboculars may include an 
additional (supplementary) scale. The anterior temporals are made up of the first and 
second row of scales. Bilateral reductions of the number of longitudinal dorsal scale 
rows (dsr, at midbody: msr) are either low (lateral) or high (i.e., paravertebral or ver- 
tebral). Their position on the posterior portion of the trunk is expressed as a percent- 
age of the total number of ventrals (%ven) and calculated on the basis of the average 
of the right and left side counts. Vertebra measurements, and in particular the length of 
the centrum (lc) and neural crest (nc), the least width of the neural arch (wn), and the 
width across the prezygapophyses (wp), are figured in Helfenberger (2001). 

Scientific names of Hierophis spp. are cited with the author and year of publi- 
cation upon their first mention in the text. In the case of Eirenis spp. except those used 
for molecular analyses (see below), as well as Oriental racer genera cited in the 
Discussion, only the author's name is given. 

Acronyms used in the text are CAS (California Academy of Sciences, San 
Francisco), MHNG (Muséum d'histoire naturelle, Genève), MVZ (Museum of Verte- 
brate Zoology, Berkeley), and ZMUC (Zoologisk Museum, K0benhavn). 



HIEROPHIS AND ZAMENIS ANDREANA 241 

Hierophis Fitzinger (sensu Utiger & Schätti, 2004) is understood to contain two 
European taxa, the endemic Cyprus whip snake, three eastern Mediterranean species, 
as well as the eastern Palaearctic H. spinalis. Lacépède's whip snake (H. viridiflavus) 
from NE Spain to Dalmatia, Malta, and Gyaros Island (Cyclades) in the Aegean Sea, 
the Balkans whip snake (H. gemonensis) from Dalmatia to the Aegean region, and H. 
cypriensis make up the western species group. Eastern Mediterranean species include 
H. caspius from the Balkans eastward and two Anatolia-Caucaso-Iranian species, H. 
jugularis and H. schmidti. As to the systematic make-up of dwarf snakes, the reader is 
referred to the appropriate section in the text. 

External morphological data for Coluber (s.l.) andreanus are from five speci- 
mens reported earlier (Schätti, 2001) and MHNG 2626.59. Scale features of Eirenis 
and Hierophis spp. are from Boulenger (1894, 1914), Eiselt (1976), Schmidtler & 
Schmidtler (1978), Doczenko (1985), Schätti (1985, 1987, 1988), Doczenko (1989), 
Franzen & Sigg (1989), Schmidtler & Eiselt (1991), Böhme (1993), Schätti (1993), 
Schmidtler (1993), Schmidtler & Baran (1993b), Schmidtler (1997), Sivan & Werner 
(2003), and Utiger & Schätti (2004). 

A partial sequence of the mitochondrial small ribosomal subunit (12S rRNA) of 
six racer or dwarf snake taxa and presumably related colubrines was obtained from 
frozen muscle or a dry skin (Eirenis sp.). Voucher specimens are Coluber (s.l.) 
andreanus (Werner, 1917) MHNG 2626.59 (Iran: Gavilah, Khuzestan), Eirenis medus 
(Chernov in Terentjev & Chernov, 1940) MHNG 2627.4 (Iran: vicinity of Mahniyah, 
Hamadan/Zanjan border), E. punctatolineatus (Boettger, 1892) MHNG 2626.99 (Iran: 
vicinity of Hashtijan, Fars), Eirenis sp. MHNG 2626.71 (Iran: Mehkuyeh, Fars; 
sloughed skin, see Dwarf Snakes), Lytorhynchus diadema (Duméril, Bibron & 
Duméril, 1854) MHNG 2427.32 (Yemen: Jabal Mafluq), and Spalerosophis microlepis 
Jan, 1865 MHNG 2626.70 (Iran: Mehkuyeh, Fars). 

The technique of DNA isolation as well as PCR and sequencing procedures are 
described in Utiger et al. (2002). The obtained data were added to a selection of 16 
Palaearctic, Saharo-Sindian, and an Oriental racer species from an existing sequence 
file consisting of two partially analysed mitochondrial genes, cytochrome oxidase sub- 
unit I (COI) and 12S rRNA (Utiger & Schätti, 2004). Due to constraints in laboratory 
facilities, only one gene region (12S rRNA) was investigated for the new taxa pre- 
sented in this study. The lacking COI characters were coded as missing. 

Phylogenetic analyses were performed with PA UP* version 4.0b 10 for Mac 
(Swofford, 1998). Weighted maximum parsimony (MP) procedures are described in 
Utiger et al. (2002) and Utiger & Schätti (2004). Gaps in the 12S rDNA sequence were 
treated as fifth character state. After a first run with heuristic search and tree-bisection 
reconnection (TBR) branch swapping, characters were weighted with the rescaled 
consistency index (RC, Farris, 1989) and a second heuristic search was performed. The 
procedure was repeated once; further weighting did not alter the parameters of the 
resulting tree. Nonparametric bootstrap values (Felsenstein, 1985) with 1000 replicates 
were calculated for unweighted and weighted characters. 



242 



B. SCHÄTTI & P. MONSCH 



Table 1. Sequence properties and tree reconstruction parameters 

gaps: missing gaps: 5 th character state 

unweighted MP unweighted MP weighted MP 



Length of sequence alignment (COI/12S) 
Total of variable characters (COI/12S) 
- parsimony-informative ((COI/12S) 
Number of most parsimonious trees 
Tree length 
Rescaled consistency index (RC) 



017 (407/610) 


1017 (407/610) 


1017 (407/610) 


339 (142/197) 


350(142/208) 


350 (142/208) 


249(116/133) 

A 


256(116/140) 

2 

1121 


256(116/140) 
i 


1067 


266.72 


0.195 


0.199 


0.478 



RESULTS 

Hierophis Fitzinger, 1834 

Loreal and preocular single (the latter rarely divided). Usually eight supra- 
labials (seven to nine), fourth and fifth entering eye. Anterior subocular situated 
between third and fourth supralabial; sometimes very small or, rarely, lacking in 
Hierophis jugularis; occasionally absent or with an additional small scale in H. 
spinalis. Two postoculars, normally only upper in contact with parietal. Usually two 
(one to, sometimes, three) scales in first row of temporals, and two or three (rarely one) 
in second. Nine or ten (eight to twelve) sublabials (Tb. 3). 

Ventrals in S S 163-174 (gemonensis) and 178-211, 9 9 168-182 and 194-227, 
respectively; subcaudals 86-125 (6 6) and 84-117 (9 9). Maximum total length ca. 
100 cm (cypriensis, gemonensis, spinalis) to over 200 cm (caspius, jugularis). 

Table 2. Selected morphological data for Hierophis spp. Dorsal scale rows (dsr) on the neck, at 
midbody, and prior to the vent, posterior reduction pattern (prp: low (1) or high (h), see Material 
and Methods), number of maxillary (max), palatine (pal), pterygoid (pter), and dentary (den) 
teeth as well as vertebrae ratios (length of the centrum: lc, length of neural crest: nc, least width 
of neural arch: wn, width across prezygapophyses: wp). 



Species 


dsr 


prp 


max 


pal 


pter 


den 


lc/wn 


lc/wp 


nc/wn 


caspius 


17-19-15 


1 - l(-h) 


12-15 


9 


14-18 


14-17 


1.25-1.44 


0.70-0.77 


0.88-1.05 


cypriensis 


17-17-13 


1-h 


16-18 


10-11 


17-20 


19-20 


1.38-1.58 


0.74-0.80 


1.11-1.28 


gemonensis 


17-19-15 


1-1 


16-19 


9-12 


20-23 


16-23 


1.23-1.35 


0.70-0.74 


0.95-1.03 


jugularis 


17-19-15 


1-1 


12-14 


11 


18-20 


16-20 


1.16-1.36 


0.71-0.73 


0.94-1.04 


schmidti 


17-19-15 


1-1 


12-15 


9 


17-19 


14-17 


1.36-1.39 


0.72-0.75 


1.00-1.08 


spinalis 


17-17-15 


1 


13-15 


8-10 


10-12 


16-18 


1.45-1.53 


0.74-0.81 


1.05-1.21 


viridiflavus 


17-19-15 


1-1 


14-16 


9-11 


15-20 


15-18 


1.28-1.44 


0.68-0.82 


0.88-1.05 



Dorsal scales with paired apical pits, in 17 longitudinal rows at neck (15 th 
ventral), 17-19 msr, and 13-15 in front of vent. A lateral increase of the number of dsr 
on anterior portion of trunk except in Hierophis cypriensis and H. spinalis. One 
(spinalis) or two lateral reductions involving rows 2-5 on both sides of posterior 
portion of body; a third fusion (rows 6-8), in most cases unilateral (i.e., 14 dsr prior to 
vent), sometimes occurs in H. caspius. H. cypriensis has 17 dsr throughout the 



HIEROPHIS AND ZAMEN1S ANDRE AN A 



243 



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B. SCHÄTTI & P. MONSCH 



forebody, a first reduction involving the third and fourth row at 61-68%ven, and a 
paravertebral or vertebral reduction (rows 6+7 or 7+8) to 13 dsr situated between 
66-79%ven (Tbs 2-3). 

Maxillary with 12-19 teeth, palatinum 8-12, pterygoid 10-12 (spinalis) and 14- 
23, and dentary 14-23. Vertebra ratios are: lc/wn 1.16-1.58, lc/wp 0.68-0.82, nc/wn 
0.88-1.28 (Tb. 2). 

Hemipenis with 3-6 distinct rows of spines; enlarged basal hook present in 
Hierophis cypriensis, H. gemonensis, and H. spinalis. Apex calyculate and bulbous in 
everted state (two longitudinal grooves in situ); borders of calyces denticulate prox- 
imally, usually smooth towards apex. 



Coluber (sensu lato) andreanus (Werner, 1917) 



PL 1 



Loreal and preocular single. Seven supralabials, third and fourth in contact with 
eye. Anterior subocular single, paired, very small, or absent (Schätti, 2001: Tb. 1; one 




Plate 1 
Coluber (sensu lato) andreanus ( 9 ) MHNG 2626.59 from Khuzestan (Gavilah), Iran. 

distinct subocular in MHNG 2626.59). Two postoculars, both (lower only partially) in 
contact with laterally extended parietal. First temporal single, coalesced with sixth 
supralabial in ZMUC R6044 (Schätti, 2001: Fig. I) 1 , usually two (one) scales in sec- 
ond temporal row. Eight sublabials (Tb. 3). 



1 Erroneously stated to be fused with the parietal (Schätti. 2001). 



HIEROPHIS AND ZAMENIS ANDREANA 



245 



Ventrals show a pronounced sexual dimorphism, i.e., 217-218 (two â 6) and 
247-269 (four ? ?); subcaudals 92-96 and 84-94, respectively. Maximum total length 
445+155 mm (â) and 586+164 mm (9) in syntypes (Werner, 1917). 

Dorsal scales with single apical pits (best visible on flanks), in 17 longitudinal 
rows on anterior portion of trunk and at midbody, and 17 (no reduction) or 15 dsr in 
front of vent (Schätti, 2001). CAS 100474 (â) with a lateral reduction involving row 
2+3 at 87%ven; MHNG 2626.59 ( $ ) has 17-15-17 (etc.) dsr between ventrals 221-246 
(89-99%ven), fluctuation between rows 2-4, and row 1+2 in the case of the last 
reduction to 15 dsr. 

Maxillary with 9-13 teeth (10 in MHNG 2626.59). Proximal portion of hemi- 
penis spinose (no further data available, vertebra ratios unknown). 




Hierophis caspius 
Hierophis schmidti 
Coluber (s.l.) andreanus 
— Hierophis jugular is 



30 



34 



95 



i- Eirenis modestus 



11 



97 



57 



100 



Eirenis medus* 

Eirenis punctatolineatus 
Eirenis sp* 

77. Hierophis gemonensis 

I Hierophis viridiflavus 

Hierophis cypriensis 



25 



78 



100 



92 



100 



■Hierophis spinalis 

-Platyceps ventromaculatus 

Platyceps collaris 

Platyceps najadum 



55 



39 



99 



— Spalerosophis cf. diadema 
• Spalerosophis microlepis ' 

Hemorrhois hippocrepis 

Hemorrhois raver gier i 

Hemerophis socotrae 



Coelognathus flavolineatus 



Lytorhynchus diadema 

10 changes 



Fig. 1 



Weighted maximum parsimony (MP) tree of two partially sequenced gene regions (COI and 12S 
RNA) with the Oriental racer Coelognathus flavolineatus (Schlegel) as outgroup. In the case of 
taxa with an asterisk reconstruction is based on 12S rDNA sequence data only. Bootstrap values 
(1000 replicates) from unweighted (upper) and weighted (lower values) MP analysis. 



246 b. schätti & p. monsch 

Comparative Morphology 

Coluber (s.l.) andreanus differs from Hierophis spp. in lower supralabial and 
sublabial counts (7 instead of 8 and 8 vs. usually 9-10, respectively), parietal shape, 
i.e., laterally extended and in contact with lower postocular, and a single first temporal 
scale (parietal usually not reaching beyond upper postocular, 2 first temporals) 2 , 247 or 
more ventrals in females (227 or less), single apical pits (paired), only one lateral dsr 
reduction that is sometimes absent (invariably present, Tb. 3), and a maximum total 
length of ca. 75 cm vs. 100-200 cm or more. Andreas' racer possesses a surprising 
number of external morphological character states in common with Eirenis spp. (see 
Tbs 3-4 and Discussion). 

Hierophis cypriensis differs from congeneric species in, for instance, the occur- 
rence of a paravertebral or vertebral reduction on the posterior portion of the trunk 
(Utiger & Schätti, 2004), and it has the highest lc/wn and nc/wn ratio within the genus 
(Tbs 2-3). 

Due to similar osteological (basisphenoid) and hemipenis features (bulbous 
shape and general ornamentation), the senior author assumed that Masticophis spinalis 
Peters from Central Asia to Korea belonged to Hierophis (Schätti, 1987: Figs 1, 3.e; 
Schätti, 1988: Figs 6, 8). H. spinalis is exceptional among Palaearctic whip snakes in 
only one reduction on the posterior portion of the trunk involving lateral scale rows, 
low tooth counts of the pterygoid (Tb. 2), and coloration, i.e., presence of a light 
vertebral stripe and no ontogenetic change of the dorsal colour pattern. 

The geographically isolated Hierophis spinalis and H. cypriensis differ from 
congeneric species in, for instance, a lower number of dsr (17 instead of 19) on the 
forebody (Tb. 2), presence of a basal hook (also in H. gemonensis, see Character 
Phylogeny), the shape of the lateral processus of the pterygoid (simple instead of bi- 
cuspid), and higher vertebra ratios (lc/wn, nc/wn, Schätti, 1987: Fig. 2). It also appears 
that female H. spinalis attain larger maximum size than males, whereas the contrary 
applies to congeneric species except H. cypriensis. 

Dwarf Snakes 

As generally understood (e.g., Doczenko, 1985; Leviton et al, 1992), dwarf 
snakes contain the putatively monotypic genus Pseudocyclophis Boettger, i.e., P. per- 
sicus (Anderson) (including, e.g., P. walteri Boettger) reported from eastern Anatolia 
to Pakistan, as well as Eirenis Jan. Eirenis africanus (Boulenger) from NE Sudan to 
Djibouti (and probably NW Somalia) as well as 16 valid species from the eastern 
Mediterranean area to the Caucasus, Iran, and Turkmenistan are recognised, i.e., E. au- 
rolineatus (Venzmer), E. barani Schmidtler, E. collaris (Ménétriés), E. coronella 
(Schlegel), E. coronelloides (Jan), E. decemlineatus (Duméril, Bibron & Duméril), E. 
eiselti Schmidtler & Schmidtler, E. hakkariensis Schmidtler & Eiselt, E. levantinus 
Schmidtler, E. lineomaculatus Schmidt, E. medus (Chernov), E. modestus (Martin), the 
type species (Schmidtler & Schmidtler, 1978), E. punctatolineatus (Boettger), E. 



2 The antero-lateral extension of the parietal, and its contact with the lower postocular, is 
correlated with the condition of the first temporal scale (single). 



HIEROPHIS AND ZAMEN1S ANDREANA 247 

rechingeri Eiselt, E. rothi (Jan), and E. thospitis Schmidtler & Lanza. Nine Eirenis spp. 
(53%) were described or revalidated (E. coronelloides) over the last 33 years. 

Pseudocyclophis persicus attains a total maximum length of ca. 50 cm and is 
characterised by the usual absence of a loreal (small if present) and a single postocular 
(Tbs 3-4). P. persicus has 15 msr and sometimes a posterior dsr reduction, 2 (1+1) 
anterior temporals, a slender habitus as well as high ventral (up to 224 in 6 S , 238 in 
9 9) and subcaudal counts (95 and 85, respectively, Doczenko, 1985). 

Doczenko (1989) assigned Eirenis collaris, E. coronella, E. eiselti, E. medus, E. 
rechingeri, and E. rothi, i.e., species with 15 msr except E. africanus (not examined), 
to a new subgenus, Collana. Schmidtler & Eiselt (1991) noted that, for instance, the 
closely related E. hakkariensis and E. thospitis with 17 and 15 msr, respectively (see 
below), caused considerable doubt ("erhebliche Bedenken") as to the taxonomic 
division of Eirenis Jan, and later studies (e.g., Schmidtler, 1993: 94) called the validity 
of Collana into question. Finally, the close relationship ("nahe Verwandtschaft") of E. 
modestus (17 msr) with E. thospitis induced Schmidtler (1997) to abandon Doczenko 's 
(1989) subgeneric concept. 

Nine of the 17 species including Eirenis coronelloides and E. thospitis have 15 
msr and there is no posterior dsr reduction except in E. coronella (fide Schmidtler & 
Schmidtler, 1978) and E. africanus (Ineich, 2003) 3 . 

Eirenis collaris, E. eiselti, E. medus, and E. rothi make up the "E. collaris 
group" (e.g., Schmidtler & Baran, 1993a) that may have evolved from E. modestus 
("Linie E. modestus - E. collaris - E. rothi", Schmidtler, 1993) or from a common 
ancestor stock (Schmidtler, 1997). A single postocular sometimes occurs in E. medus 
(Tb. 3). E. rothi often has only 7 sublabials and a single second temporal, i.e., only 2 
instead of usually 3 anterior temporals. In this species, the shape of the parietal (see 
footnote 2) is a variable feature as exemplified by MHNG 1247.1 (borders lower 
postocular) and MHNG 1363.58 (only in contact with upper postocular). 

Single second temporals are normally also found in Eirenis coronella. This 
small species (ca. 35 cm maximum total length) sometimes has 9 sublabials, a reduc- 
tion from 15 (msr) to 13 dsr in front of the vent, and low ventral (125 in â â , 143 in 
9 9) and subcaudal counts (Schmidtler & Schmidtler, 1978; Sivan & Werner, 2003). 
E. coronelloides is a minuscule (ca. 25 cm maximum total length) eastern 
Mediterranean (southern Levant) species. It has few ventrals (S 6 123-134, 9 9 140- 
155) and subcaudals (36-48 and 33-40, respectively) and differs from E. coronella (40- 
67 subcaudals) in "having a black crown, a dark ventral stripe or both" (Sivan & 
Werner, 2003). In the latter species, Schmidtler & Schmidtler (1978) recorded a mini- 
mum of 32 subcaudals for females and 38 in male specimens. 

The phylogenetic affinities of Eirenis africanus, E. coronella, E. coronelloides, 
and the singular E. rechingeri (striped, only known from the holotype) remain unclear. 
E. africanus is rare in collections; this species has 2 (1+1) anterior temporals 
(Boulenger, 1914) but, according to Scortecci (1930: Fig.), the parietal does not touch 
the lower postocular (see footnote 2). 

3 This character state could not be ascertained for Eirenis thospitis and E. coronelloides. In the 
latter species, it is supposed that there is no difference vis-à-vis E. coronella, i.e., that coro- 
nelloides has a reduction to 13 dsr prior to the vent. 



248 B. SCHÄTTI & P. MONSCH 

The "Eirenis modestus complex" (sensu Schmidtler, 1997) includes E. thospitis 
with 15 msr and five species {aurolineatus, barani, hakkariensis, levantinus, and mod- 
estus) with 17 msr and, usually, a reduction to 15 dsr prior to the vent (see footnote 3). 

The posterior reduction may be present or absent in Eirenis aurolineatus and E. 
levantinus. E. modestus has exceptionally ("sehr selten", Schmidtler & Baran, 1993b) 
19 msr; this number is more often found in E. m. cilicius Schmidtler from around 
Burdur, Turkey (Schmidtler, 1997). Specimens ofE. modestus with 17 dsr at midbody 
and in front of the vent (i.e., no reduction) are documented (e.g., Schmidtler & Eiselt, 
1991; Schmidtler, 1993: Tb. 1). 

A considerable number (33%, n=13) of northern Anatolian Eirenis m. modestus 
examined by Schmidtler (1997) have a single postocular and, rarely, there are 2 
temporals in the first row (Schmidtler & Baran, 1993b). All taxa of the "E. modestus 
complex" show variation in the number of preoculars (1-2), although two scales are 
uncommon in E. levantinus and E. hakkariensis (Schmidtler & Eiselt, 1991: Tb. 1; 
Schmidtler, 1993: Tb. I) 4 . The latter species is comparatively large (ca. 50 cm 
maximum total length) and has 8-9 sublabials. Schmidtler & Eiselt (1991) discuss 
differences of the Hakkari dwarf snake and E. punctatolineatus (see below). 

Eirenis lineomaculatus with a posterior reduction from 17 to 15 dsr involving 
lateral rows is among the smallest species (ca. 30 cm maximum total length, 
Schmidtler & Schmidtler, 1978) of the genus (see E. coronelloides). E. lineomaculatus 
is outstanding for a usually single postocular and absence of the loreal, or its small size 
if present (Tbs 3-4), as well as extremely low ventral (103-119 in S S , 117-132 in 9 9) 
and subcaudal (28-38 and 21-33, respectively) counts (Schmidtler & Schmidtler, 
1978). 

Eirenis decemlineatus and E. punctatolineatus possess a posterior reduction 
from 17 to 15 dsr involving lateral rows. The latter is remarkable for an elevated 
number of sublabials (9, sometimes 10) in the nominate subspecies (Tbs 3-4); E. p. 
kumerloevei Eiselt sometimes has 8 sublabials (Eiselt, 1976: Tb.; Franzen & Sigg, 
1989: Tb. 1). E. decemlineatus, with a total length of up to 90 cm by far the largest 
dwarf snake, has 7-9 sublabials and may have 3 second temporals. Schmidtler & Eiselt 
(1991) noted that these species are probably not closely related ("vermutlich nicht 
einmal näher miteinander verwandt"), and that the intraspecific ("geographische") 
variation of E. decemlineatus requires further studies. 

A sloughed skin of an Eirenis sp. from Fars Province (Iran, see Material and 

Methods) lacked the anteriormost portion (including head) and tail tip and could not be 

identified with certainty. Given the length of the fragment (ca. 65 cm), 17 msr, and 

clear genetic differences vis-à-vis E. punctatolineatus (Fig. 1), it cannot be excluded 

that the exuvia is from E. decemlineatus. Eiselt (1971) noted that this species had not 

yet been recorded from the vicinities of Shiraz. 

The dwarf snake fauna of Iran is not yet well explored as exemplified by Eirenis medus 
(see Material and Methods). It appears that Eiselt's (1971) reference to E. decemlineatus and the 
presence of this species in "Iran" as noted by Leviton et al. (1992) rely on Wall's (1908) "Confia 



4 Most probably, the alleged absence of a preocular in certain specimens of Eirenis barani and 
E. levantinus (Schmidtler, 1993: Tb. 1) is in error. 



HIEROPHIS AND ZAMENIS ANDREANA 249 

decemlineatcC , from Maidan Mihaftan (Masjed Soleyman, Khuzestan). The specimens having 
"In the posterior part [...] spots [that] become rearranged in longitudinal interrupted lines" most 
probably belong to E. punctatolineatus. This pattern is unknown in E. decemlineatus (F. J. 
Schmidtler in litt. August 2003). 

Character Phylogeny 

Schätti (2001) regarded the outstanding features of Coluber (s.l.) andreanus, 
i.e., the low number of supralabial, sublabials, and temporal scales, the occasional 
absence of an anterior subocular, few dsr, and the absence of posterior reductions (or a 
single fusion involving paraventral rows) as probably ancestral character states 
("wahrscheinlich plesiomorphe Merkmalszustände"). Mostly single preoculars, two 
supralabials entering the orbit, and the absence of high (paravertebral or vertebral) dsr 
reductions in most species led Schätti & Utiger (2001) consider Hierophis a conser- 
vative genus vis-à-vis other Palaearctic as well as Saharo-Sindian and Afro tropical 
racers and allied genera {Hemorrhois, Platyceps, and Spalerosophis spp.). Although 
molecular (mtDNA) data of the type species of Eirenis Jan (E. modestus) suggested 
sister group relationship of dwarf snakes to eastern Mediterranean Hierophis spp., the 
authors put forward putatively ancestral conditions of external morphology in dwarf 
snakes, in particular the absence of an anterior subocular and few supralabial and 
temporal scales. 

Schmidtler & Eiselt (1991) considered Eirenis modestus with 17 msr a very 
conservative ("sehr ursprüngliche") species. Taxa with 15 msr and including mostly 
small-sized ("eher kleinwüchsig") species were thought to be more advanced 
("fortschrittliche") dwarf snakes (Doczenko, 1989; Schmidtler, 1993: 90). New studies 
using multivariance analyses (Schmidtler, 1993, 1997) suggest that E. decemlineatus, 
the largest dwarf snake, is a very conservative and systematically isolated taxon 
("extreme Außengruppe", Schmidtler, 1993: 93). 

According to Schmidtler (1997), a large loreal ("großes Frenale") and high 
degree of fragmentation of the temporals ("hoher Zerfallgrad der Temporalia") are 
derived character states. This assertion collides with results showing that Pseudo- 
cyclophis persicus (loreal absent, 1+1 anterior temporals) is a highly advanced species 
(see Discussion) as evidenced by an extremely low reduction index (19, Schmidtler, 
1993: Tb. 3). The third presumably advanced ("wohl synapomorphe") condition within 
dwarf snakes, i.e., the contact of the first temporal with the supralabial bordering the 
orbit ("häufiger Kontakt 1. Temporale und suboculares Supralabiale", Schmidtler, 
1997: 49), is purely quantitative and highly variable. In the "Eirenis modestus 
complex", for instance, values range from 0% in E. m. modestus and certain popu- 
lations of E. m. semimaculatus (Boettger) to 39% in E. aurolineatus (Schmidtler, 1997: 
Tb. 1, character 15). 

In dwarf snakes, the loss ("Verschwinden") of the loreal (in Eirenis lineomacu- 
latus and Pseudocyclophis persicus), the number of posterior (!) temporals, and in 
particular the cranial shift of the dsr reduction from 17 to 15 are correlated with the 
reduction of total length ("reduktionsrelevante Merkmale"). Dwarfism ("Verzwer- 
gung") also affects the diameter of the eye, width of the frontal, dimensions of the 
anterior and posterior inframaxillary scales, and, in males, the number of subcaudals. 



250 B. SCHÄTTI & P. MONSCH 

The number of preocular, postocular, and gular scales are not influenced by this phe- 
nomenon (Schmidtler, 1997). 

Without anticipating the situation in Hemerophis socotrae (Günther), the gen- 
era Hemorrhois, Platyceps, and Spalerosophis, as well as the unassigned racers 
Coluber (s.l.) dorri (Lataste), C. (s.l.) scorteceli (Lanza), and C. (s.l.) zebrlnus 
(Broadley & Schätti) from the western Sahel (Sudanese Arid), Somalia, and Namibia, 
respectively (see Schätti & Utiger, 2001), the morphological and molecular data pre- 
sented in this study, and in particular the phylogenetic affinities of C. (s.l.) andreanus 
and Elrenls spp., suggest character polarities for Palaearctic whip and dwarf snakes as 
listed in Table 4. 

Table 4. Distribution of character states in dwarf and whip snakes. Taxa and their abbreviations 
are Coluber (s.l.) andreanus (A), Eirenis lineomaculatus (B), E. p. punctatolineatus (C), Eirenis 
spp. (D, see Tb. 3), Hierophis caspius, H. jugularis, and H. schmidti (E), H. gemonensis and H. 
viridiflavus (F), H. cypriensis (G), H. spinalis (H), and Pseudocyclophis persicus (I). Characters 
(0 for ancestral, 1 for derived condition) are (1) loreal: present (0), absent (1); (2) postocular(s): 
paired (0), single (1); (3) subocular: present (0), absent (1); (4) sublabials: 9 or more (0), 7 or 8 
(1); (5) supralabials: 8 or more (0), 7 (1); (6) first temporal(s): paired (0), single (1); (7) second 
temporal(s): 2 or 3 (0), single (1); (8) midbody dorsal scale rows: 19 (0), 15-17 (1); (9) posterior 
reduction(s): 2, rarely 3 (0), 1 or absent (1); (10) apical pits: paired (0), single (1). Conditions 
found in 95% or more of the individuals per species are shown; not considered are, in particular, 
the occasional occurrence of a single postocular in E. medus (Tb. 3) or rare character states found 
in other Eirenis spp., e.g., a single postocular and two first temporals (modestus) or the occasion- 
al presence of 9 sublabials {decemlineatus, hakkariensis, see Dwarf Snakes). 



no. 


character 


A 


B 


c 


D 


E 


F 


G 


H I 


1 


loreal 





1 

















1 


2 


postocular(s) 





1 

















1 


3 


anterior subocular 


0/1 


1 


1 


1 











0(1) 1 


4 


sublabials 


1 


1 


0(1) 


1 











1 


5 


supralabials 


1 


1 


1 


1 











1 


6 


first temporal(s) 


1 


1 


1 


1 











1 


7 


second temporal(s) 


0(1) 


0(1) 





0(1) 











1 


8 


midbody scale rows 


1 


1 


1 


1 








1 


1 1 


9 


posterior reduction(s) 


1 


1 


1 


1 











1 1 


10 


apical pits 


1 


1 


1 


1 











1 



Stickel (1951) and Doczenko (1985: Fig.) studied hemipenis features of Eirenis 
modestus and dentigerous bones (palatinum, pterygoid) of E. collaris, E. modestus 
auct., and Pseudocyclophis persicus. However, published information about phyloge- 
netically significant osteological (skull, vertebrae) and hemipenis characters are lack- 
ing for most dwarf snake species and Coluber (s.l.) andreanus. According to Sivan & 
Werner (2003), Shwayat (1998) investigated "hemipenis morphology" of Jordan 
Eirenis spp. 

The striking resemblance of Hierophis jugularis, H. viridiflavus, Eirenis mod- 
estus (and possibly other dwarf snake species), and Palaearctic ratsnakes as, for in- 
stance, Elaphe schrenckli Strauch and Zamenis longlssimus (Laurenti) with respect to 
hemipenis ornamentation and its bulbous shape (Schätti, 1988: Figs 6-7; Schmidtler & 
Baran, 1993b: Fig. 55; Utiger et al., 2002: Fig. 5) is most probably based on ancestral 



HIEROPHIS AND ZAMENIS ANDREANA 25 1 

character states. This is also the case with the enlarged basal spine found in Hierophis 
spp. {cypriensis, gemonensis, spinalis), E. modestus, and ratsnakes, e.g., Zamenis 
lineatus (Camerano) and species of the genera Euprepiophis Fitzinger and Oreophis 
Utiger, Helfenberger & Schätti. 

DISCUSSION 

Without taking account of the Oriental racer genera Coelognathus Fitzinger, 
Gonyosoma Wagler (including, e.g., Herpetodryas frenatus Gray and Coluber prasinus 
Blyth), and Ptyas Fitzinger (see Utiger, 2002), our molecular approach (Fig. 1) reveals 
three major evolutionary lineages of Old World colubrines. They are represented by 
five Palaearctic and Saharo-Sindian genera (Eirenis, Hemorrhois, Hierophis, 
Platyceps, Spalerò sophis), Saharo-Sindian leaf-nosed snakes {Lytorhynchus spp.), and 
the Socotra racer (Hemerophis socotrae). The uncorrected sequence divergence (p) for 
12S rDNA of H. socotrae and Lytorhynchus diadema is 14%, and, for instance, 10% in 
the case of H. socotrae and Hierophis viridiflavus. 

The geographically isolated Namibia racer Coluber (s.l.) zebrinus belongs to a lineage 
that diverged from the common racer stock prior to the evolution of Afrotropical, Palaearctic, 
and Saharo-Sindian racers and related genera (Schätti & Charvet, 2003). The sequence diver- 
gence (p) vis-à-vis Hemerophis socotrae and Hierophis viridiflavus is 14,5% and 12%, respec- 
tively (unpubl. data). 

Whip snakes of the genus Hierophis (sensu Utiger & Schätti), Coluber (s.l.) 
andreanus, and Eirenis spp. are monophyletic (Fig. 1). They represent a distinct evo- 
lutionary lineage probably including Pseudocyclophis persicus. 

MtDNA sequences corroborate preliminary molecular data (12S and COI 
rDNA) for Eirenis modestus and the phylogenetic affinities of dwarf snakes, paraphyly 
of Hierophis (sensu Utiger & Schätti), close relationship of H. cypriensis, H. gemo- 
nensis, and H. viridiflavus, and sister group status of the western whip snakes to the 
eastern Palaearctic H. spinalis (Schätti & Utiger, 2001; Utiger & Schätti, 2004). 
Furthermore, the phylogenetic tree (Fig. 1) indicates an early separation of western 
Hierophis spp. from a common ancestor stock giving rise to Palaearctic whip and 
dwarf snakes and suggests that Eirenis spp. are the sister group of Coluber (s.l.) 
andreanus and the eastern Mediterranean H. caspius, H. jugularis, and H. schmidti. 

With regard to external morphology, European and eastern Mediterranean 
Hierophis spp. show ancestral character states. Most of these species are remarkable 
for large body size. The smaller H. cypriensis and H. spinalis are advanced whip 
snakes on the basis of low msr. The absence of an increase of dsr on the forebody in 
these taxa (present in congeneric species) is correlated with the number of msr. The 
single posterior dsr reduction and, rarely, the absence of an anterior subocular in H. 
spinalis are derived conditions. 

Apart from Oriental genera (see above), Hierophis caspius and H. jugularis are 
the largest Old World racers. The reduction of body size is a remarkable evolutionary 
trend in Palaearctic whip and dwarf snakes affecting various scale characters as exem- 
plified by the small Eirenis lineomaculatus, but not the minuscule E. coronelloides (see 
Dwarf Snakes and Character Phylogeny). 

Character state distribution within the ingroup (Tb. 3) suggests that para- 
vertebral and/or vertebral dsr reductions found in Hierophis cypriensis may be derived 



252 B. SCHÄTTI & P. MONSCH 

conditions. However, the occurrence of high reductions in H. caspius (Schätti, 1988) 
and the criterion of the correlation of transformation series (Hennig, 1979) might 
favour the opposite view. In the case of the presence of a light vertebral stripe in 
H. spinalis, the outgroup criterion within whip and dwarf snakes supports auta- 
pomorphy for this character state. This might also be true for larger female maximum 
size of H. spinalis or the distinct sexual dimorphism of ventral scales in Coluber (s.l.) 
andreanus. Coloration features including melanism in Mediterranean Hierophis spp. 
are possibly correlated with large body size. 

Based on the conditions in Eirenis collaris, E. modestus, and Pseudocyclophis 
persicus (Doczenko, 1985: Fig.), the shape of the lateral processus of the pterygoid in 
Hierophis cypriensis and H. spinalis (see Comparative Morphology) may be derived 
states. The character phylogeny of other osteological conditions found in these species, 
i.e., high lc/wn and nc/wn ratio, or few pterygoid teeth in H. spinalis, are difficult to 
assess for the lack of comparative material. 

On higher systematic levels, the slow evolving 12S rDNA generally provides 
better results than the protein-coding COI sequence. Although 12S sequence data (Fig. 
1) suggest paraphyly of eastern Mediterranean Hierophis spp., we consider the generic 
allocation of Coluber (s.l.) andreanus as debatable. Reservations concerning the asso- 
ciation of Andreas' racer with these whip snakes stem from morphological evidence, 
i.e., a considerable number of derived character states supporting sister group relation- 
ship of Andreas' racer to dwarf snakes. 

Coluber (s.l.) andreanus, Eirenis spp., and Pseudocyclophis persicus have a low 
number of supralabial (third and fourth bordering eye), sublabial (except E. p. puncta- 
tolineatus), and anterior temporal scales, the parietal in contact with the lower post- 
ocular (see footnote 2), 15-17 msr with single apical pits (Boulenger 1894; Böhme, 
1993), and at most one posterior dsr reduction involving lateral or paraventral rows 
(Tbs 3-4). The occasional absence of an anterior subocular in C. (s.l.) andreanus is the 
normal condition encountered in dwarf snakes. Furthermore, the maximum total length 
of these taxa does not exceed 100 cm. 

Assuming the character phylogeny determined for whip and dwarf snakes 
(Tb. 4), an evolutionary history as indicated by molecular data (Fig. 1) would imply 
parallelism in Coluber (s.l.) andreanus and Eirenis spp., or postulate reversal of the 
polarity in at least six transformation series (sublabials, supralabials, first temporal, 
msr, dsr reduction, and apical pits) in both eastern Mediterranean whip snake sections, 
i.e., Hierophis jugulari s and H. caspius - schmidti. 

Derived conditions of head pholidosis including the occasional absence of an 
anterior subocular in Coluber (s.l.) andreanus as well as derived dorsal scale features 
(few msr, at most one posterior dsr reduction, low reduction levels, and single apical 
pits) advocate monophyly of Andreas' racer, Eirenis spp., and Pseudocyclophis per- 
sicus. The occurrence of few body scale rows (dsr, msr) and lateral (low) reductions in 
Hierophis spinalis are due to parallelism. 

Certain derived character states met with in Coluber (s.l.) andreanus, e.g., the 
absence of a dsr reduction or only 2 (1+1) anterior temporals, are characteristic for 
Eirenis africanus, E. coronella, E. rothi, and Pseudocyclophis persicus. The latter 
species, morphologically the most strongly advanced dwarf snake, and the minute E. 



HIEROPHIS AND ZAMENIS ANDREANA 253 

lineomaculatus are distinguished by the absence of a loreal and single postoculars 
(synapomorphies). Remarkably, P. persicus and C. (s.l.) andreanus are similar in, for 
instance, their slender habitus and comparatively high ventral and subcaudal counts. 

The dilemma posed by conflicting morphological and molecular data in 
Coluber (s.l.) andreanus calls for field work and further investigation in the laborato- 
ry. Certainly, paraphyly of the eastern Mediterranean Hierophis spp. and, hence, paral- 
lelism in Andreas' racer and Eirenis spp. as evidenced by mtDNA sequences cannot be 
excluded. From the molecular point of view, the systematic position of C. (s.l.) an- 
dreanus requires confirmation because a non coding sequence (12S rDNA) from a sin- 
gle specimen was analysed, i.e., a nuclear origin cannot be ruled out. 

The examination of hemipenis and osteological characters, and more genetic 
information for Coluber (s.l.) andreanus and dwarf snake species such as Eirenis 
africanus, E. coronella, E. lineomaculatus, and Pseudocyclophis persicus are crucial 
for the understanding of the perplexing results presented in this study and the evolu- 
tion of morphological characters including body size in Palaearctic whip and dwarf 
snakes. 

Taxonomic decisions with regard to Andreas' racer and the paraphyletic whip 
snake genus Hierophis (sensu Utiger and Schätti) are postponed pending further 
studies. 

ACKNOWLEDGEMENTS 

We owe our thanks to Urs Utiger (Zürich) who realised the laboratory part of 
this paper and the phylogenetic analyses of the molecular data. Theodore J. Papenfuss 
(Berkeley, MVZ) and Robert Macey (Coalnut Creek) provided a tissue sample of 
Hierophis spinalis (MVZ 211019). The field trip to Iran in spring 2003 was partially 
sponsored by a grant of the G. & A. Claraz Schenkung. Mohammad Ayatollahi, 
Department of Wildlife, and André Klaus, Swiss Embassy (both Tehran) provided 
support, logistics and invaluable help. We highly appreciated the company of our 
driver, translator, and friend Nader Rezayeh (Tehran). Josef F. Schmidtler (Munich) 
verified the determination of Eirenis medus from the Zagros and provided information 
about Iranian dwarf snakes. Corinne Charvet (Geneva) scanned and edited the illus- 
tration of Andreas' racer. Paul and Laura Joseph (London) gave linguistic advice on a 
Pacific beach. Patrick Deese and George C. Fields, Jr. (San Miguel de Allende) read 
bits and pieces of the manuscript. 

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Revue suisse de Zoologie 111 (2): 257-284; juin 2004 



Three new species of the genus Gammarus from tributaries 
of the Hi River, China (Crustacea, Amphipoda, Gammaridae) 

Zhong-e HOU 1 , Shuqiang LI 1 & Dirk PLATVOET 2 

1 Institute of Zoology, Chinese Academy of Sciences, 25 Bei-Si-Huan-Xi-Lu Str. 
Beijing 100080, P. R. China. 

(All correspondence to Shuqiang Li, E-mail: lisq@panda.ioz.ac.cn) 

2 Department of Crustacea, Zoological Museum of Amsterdam, University of 
Amsterdam, Mauritskade 57, The Netherlands. 



Three new species of the genus Gammarus from tributaries of the Hi 
River, China (Crustacea, Amphipoda, Gammaridae). - Three new 
species of the genus Gammarus are described on the base of specimens 
collected from Hi River, Xinjiang, China. Gammarus montanus sp. n. is 
characterized by uropod 3 armed with long simple setae and antenna 2 
lacking calceoli; G brevipodus sp. n. is distinguished by the peduncle of 
antenna 2 with long setae along both margins and inner ramus of uropod 3 
about one-third of outer ramus. G takesensis sp. n. differs from G brevi- 
podus sp. n. by inner ramus of uropod 3 reaching three-fourths of outer 
ramus. Distribution data on these gammarids are also presented. 

Key-words: Amphipoda - Gammarus - taxonomy - Hi River - China. 

INTRODUCTION 

The Hi River, also called " Yili" or "He" River, runs through parts of China and 
Kazakhstan. This river is 950 km long, rising from Mt Tianshan (NW Xinjiang of 
China) and flowing west across the China-Kazakhstan border, through the sandy Sary- 
Ishik-Otrau Desert, into Lake Balkhash. In its upper reaches the Hi is a mountain river; 
in the Balkhash plane it is a valley river. Before running into Lake Balkhash, it forms 
a delta of about 9000 km 2 . Hi is famous for its beauty and attracts more and more 
visitors every year. In the course of an invertebrate survey of Mt. Tianshan, a collection 
trip along the Chinese part of the Hi River was made by Dr Zhixiao Liu and students 
from Xinjiang University in July and August 2001. When checking this collection, we 
found three new gammarids, which are described in the present paper. The relationship 
between the new species and other related gammarids is discussed and their known 
distribution is presented (Map 1). 

Gammarus is one of the largest genera of epigean freshwater amphipods. 
Barnard & Barnard (1983) reviewed the freshwater Amphipoda worldwide and 117 
species were listed under the genus Gammarus. After 1983, more than 20 species have 



Manuscript accepted 12.11.2003 



258 



Z. E. HOU ET AL. 




Map 1. Localities of freshwater amphipods along the Chinese part of Hi River. Square = 
Gammarus montanus sp. n., triangle = G. takesensis sp. n., star = G. brevipodus sp. n. 



been reported by several authors (Karaman & Pinkster, 1987; Barnard & Dai, 1988 
Morino & Whitman, 1995; Stock et al, 1998; Hou & Li, 2002a, b, e, 2003a, b, e, d 
Hou, Li & Koenemann, 2002; Hou, Li & Morino, 2002; Hou, Li & Zheng, 2002 
Meng, Hou & Li, 2003). According to the study by Karaman and Pinkster (1977a, b, 
1987), these species can be subdivided into three artificial species groups, as shown in 
the key below. In the present paper, Gammarus montanus sp. n. belongs to the G. 
pulex-group, while G. brevipodus sp. n. and G. takesensis sp. n. belong to the G. bal- 
canicus-group. 



Key to species-groups in the genus Gammarus 

1 Metasome segments with middorsal process (carina) G. roeseli-group 

Metasome segments without middorsal process 2 

2 Pereopods 3 and 4 and uropod 3 bearing numerous long setae . G. pulex-group 
Pereopods 3 and 4 and uropod 3 poorly setose G. balcanicus-group 



MATERIAL AND METHODS 

Specimens were collected by a net and then preserved in 75% alcohol. For each 
species three to five specimens of each sex were dissected and appendages were 
mounted on slides according to the methods described by Holsinger (1967). The 
drawings were made with the aid of a drawing tube mounted on an Olympus BX-41 
compound microscope. 



THREE NEW GAMMARUS SPECIES FROM CHINA 259 

All holotypes treated in this study are deposited in the Institute of Zoology, 
Chinese Academy of Sciences (IZCAS), Beijing, China. Paratypes are deposited in the 
Institute of Zoology, Chinese Academy of Sciences, Beijing (IZCAS), and in the 
Muséum d'histoire naturelle, Geneva (MHNG). 

DESCRIPTIONS 

Gammarus montanus sp. n. Figs 1-5 

Material: holotype, male (IZCAS-I-A0053), Tianshan Town, Zhaosu County (43.1°N, 
81.1°E), collected by Dr Zhixiao Liu and Mr Ayiheng, August 12, 2001. Paratypes: 20 males, 20 
females and 10 juveniles (IZCAS), 9 males and 9 females (MHNG), same data as for the holo- 
type. 

Etymology: the species name "montanus" refers to the habitat of this new species. 

Diagnosis: Gammarus montanus sp. n. belongs to Gammarus pulex-group and 
is characterized by uropod 3 armed with long simple setae and by antenna 2 lacking 
calceoli. 

Description of male: body 10.2 mm in length. 

Head: eyes ovate, inferior antennal sinus deep (Fig. 1A). 

Antenna 1 (Fig. 4A): peduncular articles 1-3 in length ratio 1 : 0.59 : 0.41, with 
slender setae distally; primary flagellum twenty-eight articulate, most articles with 
aesthetascs, accessory flagellum four articulate. 

Antenna 2 (Fig. 4B): peduncular article 4 about as long as article 5, both with 
two to three groups of setae along anterior and posterior margins; flagellum ten arti- 
culate, calceoli lacking. 

Upper lip convex, with minute setules (Fig. IG). 

Mandibles (Figs 11, K): left incisor with five teeth, lacinia mobilis with four 
dentitions, molar with one long plumose seta; article 2 of palp with thirteen long stiff 
slender setae on medial margin, article 3 reaching 83% length of article 2, bearing two 
groups of long B-setae on medial surface, three long A-setae on lateral surface, a row 
of eighteen short plumose D-setae and four long E-setae. Right incisor with four teeth, 
lacinia mobilis bifurcate, molar with one long plumose seta. 

Lower lip (Fig. IH): inner lobe absent. 

Maxilla 1 asymmetrical (Figs 1J, L): medial margin of inner plate bearing 
several slender setules basally, and fifteen plumose setae evenly distributed between 
the base and apex; outer plate sparsely setose medially, the apex bearing eleven serrate 
robust setae; palp two articulate, the left second article falcate, bearing eight naked 
robust setae apically accompanied by three stiff setules apico-facially; the right second 
article bearing five broad-based tooth setae and two stiff slender setae. 

Maxilla 2 (Fig. IF): inner plate with an oblique row of thirteen plumose setae 
on medial surface, evenly distributed between the base and apex; outer plate with stiff 
setae apically. 

Maxilliped (Fig. IM): inner plate bearing one subapical and three apical blade- 
like tooth setae, associated with plumose setae on medial and apical margins; outer 
plate bearing sixteen blade-like robust setae on medial margin and eight apical pecti- 
nate setae; palp four articulate. 



260 



Z. E. HOU ET AL. 




FlG. 1. Gammarus montantes sp. n., holotype, maie. A, head; B, epimeral plate 1; C, epimeral 
plate 2; D, epimeral plate 3; E, urosomites (dorsal view); F, maxilla 2; G, upper lip; H, lower lip; 
I, right mandible; J, palp of right maxilla 1; K, left mandible; L, left maxilla 1; M, maxilliped. 



THREE NEW GAMMARUS SPECIES FROM CHINA 



261 




Fig. 2. Gammarus montanus sp. n. holotype, male. A, propodus and dactylus of gnathopod 1; B, 
propodus and dactylus of gnathopod 2; C, gnathopod 2; D, gnathopod 1. 



262 Z. E. HOU ET AL. 

Coxal plates: coxal plates 1-3 subrectangular (Figs 2C, D, 3E), bearing two to 
three seniles on anterior corner and one setule on posterior corner; coxal plate 4 exca- 
vate (Fig. 3A), bearing two setules on anterior corner and six setules on posterior 
margin; anterior lobe of coxal plates 5 and 6 small (Figs 3B, C), posterior lobe with 
two setules on posterodistal corners; coxal plate 7 (Fig. 3D) with six setules on 
posterior margin. 

Coxal gills: coxal gills present on pereopods 2-7, sac-like. 

Gnathopod 1 (Figs 2 A, D): basis with long naked setae along lateroproximal 
and medial margins, four plumose setae on posterodistal corner; carpus reaching 73% 
length of propodus, bearing two groups of long setae on lateral margin, and a row of 
long slender setae on medial margin; palm of propodus strongly oblique, bearing one 
palmar medial robust seta, ten robust setae on lateral margin and six robust setae on 
medial surface, associated with groups of long naked setae on medial surface; dactylus 
with one long naked seta on lateral margin. 

Gnathopod 2 (Figs 2B, C) larger than gnathopod 1, basis similar to that of 
gnathopod 1; carpus reaching 75% length of propodus, bearing one group of long 
naked setae on lateral margin and a row of long setae on medial margin; propodus sub- 
rectangular, bearing one blunt palmar medial robust seta, two robust setae on medial 
posterodistal corner and two robust setae on lateral posterodistal corner, with eight 
groups of long setae on medial surface; dactylus with one naked seta on lateral margin. 

Pereopod 3 (Figs 3E, F) longer than pereopod 4, basis with fourteen slender 
setae on medial margin, three groups of long setae and three groups of short setae on 
lateral margin; merus to propodus densely with long setae on medial margins; carpus 
and propodus accompanied by small robust setae; dactylus with one plumose seta on 
lateral margin and two stiff setae at hinge of nail. 

Pereopod 4 (Figs 3A, G): subequal to pereopod 3, medial margins of merus and 
carpus with less long straight setae than pereopod 3. 

Pereopod 5 (Figs 3B, H): basis nearly straight on medial margin, bearing four 
short robust seta on lateral margin and a row of eight setules on medial margin; merus 
bearing three groups of slender setae on lateral margin; carpus with two and three 
groups of short robust seta along medial and lateral margins; dactylus with 1 plumose 
seta on lateral margin. 

Pereopod 6 (Figs 3C, I): basis weakly concave posteriorly, bearing a group of 
posterodistal setae on medial surface; merus to propodus bearing two or three groups 
of robust setae on lateral margins; dactylus with one seta on lateral margin and one stiff 
seta at hinge of nail. 

Pereopod 7 (Figs 3D, J): similar to pereopod 6, basis weakly expanded posteri- 
orly, with one group of posterodistal setae on medial surface. 

Epimeral plates: epimeral plates 1-3 bearing three to four short setules on 
medial margins. Epimeral plate 1 (Fig. IB) with thirteen long setae on anteroventral 
corner and one robust seta on medial-ventral margin; epimeral plates 2 and 3 slightly 
pointed on posterodistal corners (Figs IC, D), with three robust setae and two setae on 
ventral margin of plate 2 and four robust setae on ventral margin of plate 3. 

Pleopods: pleopods 1-3 subequal in length (Figs 4F-H), peduncle bearing two 
retinacula accompanied by two setae on anterodistal corner and some long setae on 



THREE NEW GAMMARUS SPECIES FROM CHINA 



263 




Fig. 3. Gammarus montanus sp. n., holotype, male. A, pereopod 4; B, pereopod 5; C, pereopod 
6; D, pereopod 7; E, pereopod 3; F, dactylus of pereopod 3; G, dactylus of pereopod 4; H, dac- 
tylus of pereopod 5; I, dactylus of pereopod 6; J, dactylus of pereopod 7. 



264 Z. E. HOU ET AL. 

medial surface; both rami thirteen to eighteen articulate, armed with long plumose 
setae. 

Urosomites: urosomites 1-3 without humps dorsally (Fig. IE), urosomites 1-2 
bearing four groups of one or two robust setae accompanied by fine setae on postero- 
dorsal margins; urosomite 3 with two single robust setae and two single fine setae on 
posterodorsal margin. 

Uropod 1 (Fig. 4E): peduncle with one basofacial robust seta, three robust setae 
on lateral margin, one robust seta on medial margin, two robust setae on laterodistal 
corner and one robust seta on mediodistal corner; outer ramus with two robust setae on 
medial margin and two robust setae on lateral margin; inner ramus with one robust seta 
on lateral margin and two robust setae on medial margin; both rami with five apical 
robust setae. 

Uropod 2 (Fig. 4D): peduncle longer than both rami, bearing three slender 
robust setae on lateral margin, one robust seta on medial margin and one distal robust 
seta on medial and lateral corners; outer ramus a little shorter than inner ramus, bearing 
one robust seta on medial margin and one robust seta on lateral margin; inner ramus 
bearing two robust setae on medial margin and one robust seta on lateral margin; both 
rami bearing five apical robust setae. 

Uropod 3 (Fig. 41): peduncle short, bearing a pair of submarginal robust setae 
mid-laterally, one robust seta apico-laterally, two robust setae apico-medially, and two 
robust setae and a seta on the mid-vontrodistal margin; inner ramus short, about 47% 
length of outer ramus, bearing medially ten long simple setae, without lateral setae, the 
apex bearing one robust seta accompanied by five long setae; outer ramus two articu- 
late, the second article tiny, about one twentieth the length of the first, and subequal to 
the three robust apical setae of that article, the second article bearing four long apical 
setae; the medial margin of the first article with groups of long simple setae, the lateral 
margin with four robust setae associated with many long simple setae. 

Telson (Fig. 4C) cleft to basis, each lobe with two or three distal robust setae 
accompanied by two to four long slender setae, and with two groups of setae on dorsal 
surface. 

Description of female: length 7.5 mm. Propodus of gnathopod 1 ovate (Fig. 
5B), palm not oblique as in male, bearing eight robust setae on posterior margin 
accompanied by many long setae, and seven groups of setae on medial surface; nail of 
dactylus elongate, bearing one seta on lateral margin. Propodus of gnathopod 2 
subrectangular (Fig. 5A), bearing two robust setae on posteromedial corner and two 
robust setae on posterolateral corner. Uropod 3 similar to that of male, inner ramus 
reaching 45% length of article 1 of outer ramus, both rami armed with long simple 
setae (Fig. 5C). Telson cleft to the basis (Fig. 5E), each lobe with three distal robust 
setae and one basolateral robust seta. Oostegites of pereopods 2-5 present (Fig. 5D). 

Remarks: Gammarus montanus sp. n. bears characters in common with G. gre- 
goryi Tattersall, 1924 and G. paucispinus Hou & Li, 2002b particularly in the presence 
of simple setae on both margins of uropod 3, and in the shape of the epimeral plates 
1-3. G. montanus sp. n. can be distinguished from G. gregoryi by the absence of 
calceoli and the inner ramus of uropod 3 which is about half as long as article 1 of the 



THREE NEW GAMMARUS SPECIES FROM CHINA 



265 




Fig. 4. Gammarus montanus sp. n., holotype, male. A, antenna 1; B, antenna 2; C, telson; D, 
uropod 2; E, uropod 1; F, pleopod 1; G, pleopod 2; H, pleopod 3; I, uropod 3. 



266 



Z. E. HOU ET AL. 




Fig. 5. Gammarus monîanus sp. n., female. A, propodus and dactylus of gnathopod 2; B, propo- 
dus and dactylus of gnathopod 1; C, uropod 3; D, oostegite 2; E, telson. 



THREE NEW GAMMARUS SPECIES FROM CHINA 267 

outer ramus (about one-third in G. gregoryi). G. montanus sp. n. differs from G. pauci- 
spinus in that urosomite 1 bears four singly arranged robust setae on the dorsal margin 
(urosomite 1 only with a few setae in G. paucispinus), and the telson bears few long 
setae on the dorsal margin (telson with groups of long setae on dorsal margin in G. 
paucispinus). 

Gammarus brevipodus sp. n. Figs 6-10 

Material: holotype, male (IZCAS-I-A0055), from a feeder stream of the Hi River at the 
crossroad of National Highway 217 and 218, Xinyuan County (43.25°N, 83.16°E), collected by 
Dr Zhixiao Liu and Mr Ayiheng, August 16, 2001. Paratypes: 14 males and 5 females (IZCAS), 
same data as for the holotype; 10 males and 10 females (MHNG), Kunes River, Narat Town 
(43.2°N, 84.6°E), Xinyuan County, collected by Dr Zhixiao Liu and Mr Ayiheng, August 15, 
2001. 

Other material: 20 males, 9 females and 15 juveniles, Kunes River, Narat Town, 
collected by Dr Zhixiao Liu and Mr Ayiheng; August 15, 2001. 25 males, 10 females and 2 
juveniles, Bayanbulak (43.0°N, 84.1°E), collected by Dr Zhixiao Liu and Mr Meng Kaibayier, 
August 17, 2001. 

Etymology: the epithet "brevipodus" refers to the short inner ramus of uropod 3. 

Diagnosis: Gammarus brevipodus sp. n. belongs to the G. balcanicus-growp. 
Gammarus brevipodus sp. n. can be distinguished by the following characters: (1) 
antenna 2 with long setae along both margins, (2) calceoli absent, (3) uropod 3 armed 
with few setae, (4) article 2 of outer ramus in uropod 3 subequal to adjacent robust 
setae. 

Description of male: body length 14.5 mm. 

Head (Fig. 6A): lateral cephalic lobe truncate, inferior antennal sinus deep, eyes 
relatively small. 

Antenna 1 (Fig. 9F): peduncular articles 1-3 in length ratio 1 : 0.7 : 0.44, bearing 
short distal setae; primary flagellum twenty articulate, most articles with aesthetascs; 
accessory flagellum four articulate. 

Antenna 2 (Fig. 9E): peduncular article 4 a little shorter than article 5, both 
articles with long setae along anterior and posterior margins; flagellum eight articulate, 
with some distal short setae, calceoli lacking. 

Upper lip convex (Fig. 6D), with minute setules. 

Mandibles (Figs 6H, I): left incisor with five teeth; lacinia mobilis with four 
dentitions; molar triturative, bearing one plumose seta; article 2 of palp with seventeen 
long stiff setae on medial margin, article 3 reaching 80% length of article 2, bearing 
two groups of long simple B-setae on medial surface, seven long simple A-setae on 
lateral surface, a row of D-setae on medial margin and four E-setae. Right incisor with 
four teeth, lacinia mobilis bifurcate, with several weak teeth at ridge. 

Lower lip (Fig. 6C): inner lobe absent. 

Maxilla 1 (Figs 6F, G): inner plate triangular, medial margin bearing several 
slender setules basally, and twelve plumose setae evenly distributed between the base 
and apex; outer plate subrectangular, sparsely setose medially, the apex bearing eleven 
serrate robust setae; palp 2 articulate, the left second article weakly falcate, bearing 
mediodistally a single short stout seta, apically 4 robust naked setae and three stiff 
setules apico-facilly; the right second article bearing five broad-based tooth setae and 
two stiff slender setae. 



268 Z. E. HOU ET AL. 

Maxilla 2 (Fig. 6E): inner plate with an oblique row often plumose setae on the 
medial surface, evenly distributed between the base and apex; the outer plate bearing 
slender setae apically. 

Maxilliped (Fig. 6J): inner plate bearing one subapical and three apical robust 
tooth setae, associated with plumose setae on medial and apical margins; outer plate 
bearing thirteen blade-like robust setae on medial margin and 6 apical pectinate setae; 
palp 4 articulate. 

Coxal plates: coxal plates 1-3 subrectangular (Figs 7A, B, 8A), with three to 
five short setules on anteroventral corner and one seta on posteroventral corner; coxal 
plate 4 excavate (Fig. 8B), with four setules on anteroventral corner, 8 setules on 
medial margin and some minute facial setae; anterior lobe of coxal plates 5 and 6 small 
(Figs 8C, D), with one or two setules on anteroventral corner, posterior margin with 
three or two setules; coxal plate 7 (Fig. 9 A) with five setules on medial margin. 

Gnathopod 1 (Figs 7 A, C): basis with long setae on lateroproximal and medial 
margin, with four plumose setae on posterodistal corner accompanied by three long 
setae; carpus reaching 75% length of propodus, bearing one group of long setae on 
lateral margin and several groups of setae on medial margin; propodus pyriform, palm 
oblique, bearing one medial stout robust seta, ten robust setae on lateral margin and six 
robust setae on medial surface, associated with groups of long setae on medial surface; 
dactylus with 1 seta on lateral margin. 

Gnathopod 2 (Figs 7B, D): basis similar to that of gnathopod 1 ; carpus reaching 
86% length of propodus, with subparallel margins; propodus subrectangular, palm with 
one medial stout robust seta, four robust setae on lateral posterodistal corner, two 
robust setae on medial posterodistal corner, and seven groups of long setae on medial 
surface; dactylus with one seta on lateral margin. 

Pereopod 3 (Figs 8A, E) slender, basis with four long setae on lateral margin 
and fifteen setae on medial margin; merus with two single robust setae on lateral 
margin and three groups of short setae on medial margin; carpus with three short robust 
setae accompanied by short setae on medial margin; propodus with four groups of short 
robust setae on medial margin; dactylus with one seta on lateral margin and two stiff 
setae at hinge of nail. 

Pereopod 4 (Figs 8B, F): armature similar to that of pereopod 3. 

Pereopod 5 (Figs 8C, G): basis nearly straight on medial margin, with five 
single robust setae on lateral margin and a row of twenty short setae on medial margin; 
merus with two groups of short setae on lateral margin; carpus with two groups of short 
robust setae on lateral and medial margins; propodus with four groups of short robust 
setae on lateral margin and a few short setae on medial margin; dactylus with one 
plumose seta on lateral margin and 2 setae at hinge of nail. 

Pereopod 6 (Figs 8D, H): similar to pereopod 5, basis weakly concave on 
medial margin, lateral margins of merus to propodus with two to four groups of robust 
setae and fine setae. 

Pereopod 7 (Figs 9 A, G): basis weakly expanded on medial margin, medial 
surface with one robust seta on posterodistal corner. 

Epimeral plates: epimeral plates 1-3 with five to six short setules on postero- 
medial margins. Epimeral plate 1 ventrally rounded (Fig. 6L), with seven setae on 



THREE NEW GAMMARUS SPECIES FROM CHINA 



269 




Fig. 6. Gammarus brevipodus sp. n., holotype, male. A, Head; B, urosomites (dorsal view); C, 
lower lip; D, upper lip; E, maxilla 2; F, left maxilla 1 ; G, palp of right maxilla 1 ; H, left mandible; 
I, right mandible; J, maxilliped; K, epimeral plate 2; L, epimeral plate 1; M, epimeral plate 3. 



270 



Z. E. HOU ETAL. 




Fig. 7. Gammarus brevipodus sp. n., holotype, maie. A, gnathopod 1; B, gnathopod 2; C, propo- 
dus and dactylus of gnathopod 1 ; D, propodus and dactylus of gnathopod 2. 



THREE NEW GAMMARUS SPECIES FROM CHINA 



271 




Fig. 8. Gammarus brevipodus sp. n., holotype, male. A, pereopod 3; B, pereopod 4; C, pereopod 
5; D, pereopod 6; E, dactylus of pereopod 3; F, dactylus of pereopod 4; G, dactylus of pereopod 
5; H, dactylus of pereopod 6. 



272 Z. E. HOU ET AL. 

anterior corner; epimeral plate 2 with blunt posterior corner (Fig. 6K), bearing two 
ventrofacial robust setae; epimeral plate 3 with weakly pointed posterodistal corner 
(Fig. 6M), bearing one seta and two robust setae on ventrofacial margin. 

Pleopods: pleopods 1-3 subequal in length (Figs 9H-J), peduncle with two 
retinacula accompanied by two setae on anterodistal corners; both rami subequal, 
twelve to eighteen articulate, armed with long plumose setae. 

Urosomites: urosomites 1-3 without humps dorsally, urosomites 1-2 with three 
groups of robust setae accompanied by fine setae on posterodorsal margins; urosomite 
3 with two pairs of robust setae accompanied by fine setae and two fine medial setules 
on posterodorsal margin (Fig. 6B). 

Uropod 1 (Fig. 9L): peduncle longer than rami, with one basofacial robust seta, 
three robust setae on lateral margin, three robust setae on medial margin, two robust 
setae on laterodistal corner and one mediodistal robust seta; inner ramus with two 
robust setae on medial margin and one robust seta on lateral margin; outer ramus with 
two robust setae on lateral margin and one robust seta on medial margin; both rami 
with five distal robust setae. 

Uropod 2 (Fig. 9K): peduncle longer than both rami, bearing two robust setae 
on lateral margin, one robust seta on medial margin, and one robust seta on laterodistal 
and mediodistal corners; outer ramus with one robust seta on lateral margin; inner 
ramus with one robust seta on lateral margin and one robust seta on medial margin. 

Uropod 3 (Fig. 10A): peduncle short, bearing a pair of submarginal short robust 
setae mid-laterally, two robust setae apico-laterally, three robust setae apico-medially, 
and a pair of slender setules and a single robust seta on the mid-ventrodistal margin; 
inner ramus short, about one third the length of the outer ramus, bearing medially two 
robust setae, each with an associated fine seta, and a single small seta distally, without 
lateral setae, the apex bearing two robust setae, one long and two short fine setae; the 
outer ramus two articulate, the second article tiny, about one fifteenth the length of the 
first, and subequal to the six robust apical setae of that article, the second article 
bearing three fine apical setae; the medial margin of the first article with five transverse 
rows of mixed robust and fine setae, the lateral margin with five rows. 

Telson (Fig. 9B) cleft 75%, each lobe with three robust setae and two or three 
slender setae on the distal margin of each lobe, only a single facical setule on the left 
lobe. 

Description of female: body length 13.8 mm. Gnathopod 1: coxal plate with 
four setae on anterior corner and three setae on posterior corner; propodus not oblique 
as in male, with six robust setae on posterior corner (Fig. IOC). Gnathopod 2: carpus 
about as long as propodus, propodus subrectangular, with one robust seta on posterior 
corner, nail of dactylus elongate (Fig. 10D). Bases of pereopods 5-7 similar to those of 
male (Figs 10E-G). Uropod 3 (Fig. 10B): inner ramus less than 30% length of article 
1 of outer ramus. Oostegites present on pereopods 2-5 (Fig. 10G). 

Remarks: Gammarus brevipodus sp. n. is similar to G. balcanicus Schaferna, 
1923 in pereopods 3 and 4 with few setae and the shape of epimeral plates. Gammarus 
brevipodus sp. n. differs from G. balcanicus by (1) peduncle of antenna 2 with long 
setae along both margins and calceoli absent, (2) inner ramus of uropod 3 about one- 



THREE NEW GAMMARUS SPECIES FROM CHINA 



273 




Fig. 9. Gammarus brevipodus sp. n., holotype, male. A, pereopod 7; B, telson; C, terminal article 
of uropod 3; D, flagellum of antenna 1; E, antenna 2; F, antenna 1; G, dactylus of pereopod 7; 
H, pleopod 1; I, pleopod 2; J, pleopod 3; K, uropod 2; L, uropod 1. 



274 



Z. E. HOU ET AL. 




FlG. 10. Gammarus brevipodus sp. n., male: A, female: B - G. A, uropod 3; B, uropod 3; C, 
propodus and dactylus of gnathopod 1; D, propodus and dactylus of gnathopod 2; E, basis of 
pereopod 6; F, basis of pereopod 7; G, basis of pereopod 5. 



THREE NEW GAMMARUS SPECIES FROM CHINA 275 

third of outer ramus, article 2 of outer ramus of uropod 3 very short, and both rami with 
few setae in both margins, (3) telson with very few setae on dorsal margin. G. bre- 
vipodus sp. n. differs from G. montanus by pereopods 3 and 4 and uropod 3 with few 
long setae. 

Gammarus takesensis sp. n. Figs 11-15 

Material: holotype, male (IZCAS-I-A0057), Takes River (a feeder stream of the Hi 
River) near Takes County (43.13°N, 81.49°E), collected by Dr Zhixiao Liu and Mr Ayiheng, 
August 11, 2001. Paratypes: 20 males and 10 females (IZCAS), 9 males and 9 females (MHNG), 
same data as for the holotype. 

Etymology: the specific name refers to the county of origin. 

Diagnosis: Gammarus takesensis sp. n. belongs to the G. balcanicus-group and 
is characterized by: peduncle of antenna 2 with a few long setae, article 2 of outer 
ramus of urpod 3 elongate, medial margin of outer ramus and both margins of inner 
ramus of uropod 3 with short plumose setae, respectively. 

Description of male: body length 12.5 mm. 

Head: inferior antennal sinus deep, eyes medium in size (Fig. IIA). 

Antenna 1 (Figs 14B, C): peduncular articles 1-3 in length ratio 1 : 0.65 : 0.37, 
with some distal fine setae; primary flagellum twenty-eight articulate, most articles 
with aesthetascs; accessory flagellum four articulate. 

Antenna 2 (Figs 14 A, D): peduncular article 4 about as long as article 5, both 
with some short setae along anterior and posterior margins; flagellum twelve articulate, 
bearing short setae, some articles with calceoli. 

Upper lip (Fig. 11C) convex, with minute setules apically. 

Mandibles (Figs 11H, I): left incisor with five teeth; lacinia mobilis with four 
dentitions; molar triturati ve; article 2 of palp with thirteen long stiff setae on medial 
margin, article 3 reaching 75% length of article 2, with six A-setae on lateral surface 
and five long B-setae on medial surface, a row of twenty short plumose D-setae and 
five long E-setae. Incisor of right mandible with four teeth; lacinia mobilis bifurcate, 
with weak teeth at ridge; molar with one long seta. 

Lower lip: inner lobe absent (Fig. HE). 

Maxilla 1 (Figs 11F, G): inner plate triangular, medial margin bearing several 
setules basally, and fourteen plumose setae evenly distributed between the base and 
apex; outer plate subrectangular, the apex bearing eleven serrate robust setae; palp two 
articulate, the left second article weakly falcate, bearing apically seven robust naked 
setae and three stiff setules apico-facially; the right second article apically bearing four 
blunt robust setae and two stiff setae. 

Maxilla 2 (Fig. 1 ID): inner plate bearing a row of thirteen plumose setae on me- 
dial surface, evenly distributed between the base and apex; outer plate bearing long 
stiff setae apically and fine pubescence on lateral margin. 

Maxilliped (Fig. 11 J): inner plate with one subapical robust seta and three blunt 
apical robust setae, associated with plumose setae on medial and apical margins; outer 
plate with seven slender robust setae on medial margin accompanied by short naked 
setae on medial margin and six pectinate setae apically; palp four articulate. 



276 



Z. E. HOU £7 AL. 




Fig. 11. Gammarus takesensis sp. n., holotype, male. A, head; B, urosomites (dorsal view); C, 
upper lip; D, maxilla 2; E, lower lip; F, left maxilla 1; G, palp of right maxilla 1; H, right 
mandible; I, left mandible; J, maxilliped. 



THREE NEW GAMMARUS SPECIES FROM CHINA 277 

Coxal plates: coxal plates 1-3 subrectangular (Figs 12A, B, 13B), with two to 
four setules on anterior comer and one setule on posterior corner; coxal plate 4 exca- 
vate (Fig. 13E), with two setules on anterior corner and six setules on medial margin; 
anterior lobe of coxal plates 5 and 6 (Figs 13C, D) small, with one setule, posterior lobe 
with one or two setules; coxal plate 7 shallow (Fig. 13F), with three setules on medial 
margin. 

Gnathopod 1 (Figs. 12A, C): basis stout, bearing long setae on lateroproximal 
and medial margins; carpus reaching 72% length of propodus; palm of propodus 
oblique, bearing one palmar medial robust seta, five groups of robust setae on lateral 
margin, three groups of robust setae on medial surface, associated with five to seven 
groups of setae on medial surface; nail of dactylus short, bearing one naked seta on 
lateral margin. 

Gnathopod 2 (Figs 12B, D): basis similar to that of gnathopod 1; carpus 
reaching 80% of propodus, with parallel-sided margins, bearing one group setae on 
lateral margin and many setae on medial margin; propodus subrectangular, bearing one 
medial palmar robust seta, two robust setae on medial posterodistal corner, two robust 
setae on lateral posterodistal corner and seven groups of long setae on medial surface; 
dactylus with one seta on lateral margin. 

Pereopod 3 (Figs 13B, G): basis with four groups of long setae on medial 
margin; merus with one seta and one robust seta accompanied by one seta on lateral 
margin, and three groups of setae on medial margin; carpus with two pairs of robust 
setae accompanied by fine setae on medial margin; propodus with three single robust 
setae and a pair of robust setae on medial margin; dactylus with one plumose seta on 
lateral margin and two stiff setae at hinge of nail. 

Pereopod 4 (Figs 13E, H): subequal to pereopod 3, merus and carpus with less 
setae on medial margin. 

Pereopod 5 (Figs 13C, I): basis nearly straight on medial margin, bearing four 
single short robust setae on lateral margin and a row of ten short setules on medial mar- 
gin; merus bearing two single robust setae accompanied by one fine setule on lateral 
margin, and one short robust seta on medial margin; carpus with a pair of short robust 
setae on lateral margin and a group of three short robust setae on medial margin; propo- 
dus with three pair of short robust setae on lateral margin; dactylus with one plumose 
seta on lateral margin and two stiff setae at hinge of nail. 

Pereopod 6 (Figs 13D, J): similar to pereopod 5 except for basis weakly 
concave on posterior margin. 

Pereopod 7 (Figs 13F, K): similar to pereopod 5, basis slightly expanded on 
medial margin, bearing two posterodistal setae on medial surface. 

Epimeral plates: epimeral plates 1-3 with two or three short setules on postero- 
medial margins. Epimeral plate 1 ventrally rounded (Fig. 15D), bearing ten setae on 
anteroventral corner; epimeral plates 2 and 3 with slightly pointed posterior corners 
(Figs 15E, F), bearing one to two setae and two to four robust setae on ventral margin. 

Pleopods: pleopods 1-3 subequal in length (Figs 14H-J), peduncle with some 
setae on medial surface, bearing two retinacula accompanied by two setae on 
anterodistal corner; both rami eighteen to twenty articulate, fringed with plumose 
setae. 



278 



Z. E. HOU ET AL. 




FlG. 12. Gammarus takesensis sp. n., holotype, maie. A, gnathopod 1; B, gnathopod 2; C, pro- 
podus and dactylus of gnathopod 1 ; D, propodus and dactylus of gnathopod 2. 



THREE NEW GAMMARUS SPECIES FROM CHINA 



279 




Fig. 13. Gammarus takesensis sp. n., holotype, male. A, uropod 3; B, pereopod 3; C, pereopod 
5; D, pereopod 6; E, pereopod 4; F, pereopod 7; G, dactylus of pereopod 3; H, dactylus of 
pereopod 4; I, dactylus of pereopod 5; J, dactylus of pereopod 6; K, dactylus of pereopod 7. 



280 



Z. E. HOU ET AL. 




FlG. 14. Gammarus takesensis sp. n., holotype, male. A, antenna 2; B, antenna 1; C, flagellum of 
antenna 1; D, flagellum of antenna 2; E, uropod 2; F, uropod 1; G, telson; H, pleopod 1; I, 
pleopod 2; J, pleopod 3. 



THREE NEW GAMMARUS SPECIES FROM CHINA 



281 




Fig. 15. Gammarus takesensis sp. n., female: A - C, male: D - F. A, propodus and dactylus of 
gnathopod 2; B, propodus and dactylus of gnathopod 1; C, oostegite 2; D, epimeral plate 1; E, 
epimeral plate 2; F, epimeral plate 3. 



282 Z. E. HOU ET AL. 

Urosomites (Fig. IIB): urosomites 1-3 dorsally flat, bearing four groups of one 
to three robust setae accompanied by fine setae on posterodorsal margins. 

Uropod 1 (Fig. 14F): peduncle longer than both rami, bearing one basofacial 
robust seta, two robust setae on lateral margin, one short robust seta on medial margin, 
two robust setae on laterodistal corner and one robust seta on mediodistal corner; outer 
ramus with one robust seta on lateral and medial margins; inner ramus with one robust 
seta on medial margin; both rami with five distal robust setae. 

Uropod 2 (Fig. 14E): peduncle longer than both rami, bearing two robust setae 
on lateral margin, one robust seta on medial margin, and one robust seta on laterodistal 
and mediodistal corner; outer ramus shorter than inner ramus, with one robust seta on 
lateral margin; inner ramus with two robust setae on medial margin; both rami with five 
distal robust setae. 

Uropod 3 (Fig. 13 A): peduncle short, bearing a short robust seta mid-laterally, 
a single roubust seta apico-medially, and three robust setae on the mid-vontrodistal 
margin; inner ramus reaching 74% length of article 1 of outer ramus, bearing medially 
four robust setae accompanied by a row of short plumose setae, the apex bearing two 
robust setae and three long naked setae; the outer ramus two articulate, the second 
article reaching 13% length of first article, and longer than the three robust apical setae 
ofthat article, the second article bearing one fine setule apically; the medial margin of 
the first article with a row of short plumose setae and the lateral margin with three 
groups of robust setae accompanied by fine setae. 

Telson deeply cleft (Fig. 14G), each lobe with three distal robust setae accom- 
panied by one to three long setae, and 1 lateral seta. 

Description of female: body length 11.2 mm. Propodus of gnathopod 1 (Fig. 
15B) so strongly as in the male, bearing twelve robust setae on medial margin; dacty- 
lus with one seta on lateral margin, nail elongate. Propodus of gnathopod 2 with only 
slightly oblique distal margin (Fig. 15A), bearing two robust setae on medial margin. 
Oostegites of pereopods 2-5 present. 

Remarks: Gammarus takesensis sp. n. is similar to G. brevipodus sp. n. in the 
shape of gnathopods 1 and 2, and in the armature of pereopods 3 and 4 and telson. G. 
takesensis sp. n. differs from the latter by the presence of calceoli, inner ramus of 
uropod 3 reaching three-fourths of outer ramus, article 2 of outer ramus of uropod 3 
longer than adjacent robust setae, and medial margin of outer ramus and both margins 
of inner ramus of uropod 3 with short plumose setae. 

G. takesensis sp. n. is also similar to G. bosniacus Schaferna, 1923 in pereopods 
3 and 4 with few setae, and uropod 3 with some short plumose setae in medial margin 
of outer ramus. G. takesensis sp. n. differs from the latter by accessory flagellum of 
antenna 1 with four segments, antenna 2 with calceoli, and epimeral plates 2 and 3 
without long setae on ventral margins. 

ACKNOWLEDGEMENTS 

The authors would like to express their thanks to Dr Zhixiao Liu, Mr Meng 
Kaibayier and Mr Ayiheng for the donation of specimens used in this study. 



THREE NEW GAMMARUS SPECIES FROM CHINA 283 

The present study was supported by the National Natural Sciences Foundation 
of China (NSFC-30270183, 30370263, 30310464), and by the National Science Fund 
for Fostering Talents in Basic Research (NSFC-J0030092). Additional support was 
received from the Knowledge Innovation Program of the Chinese Academy of Science 
(KSCX2-1-06A, KSCX3-IOZ-01) and from the Life Science Special Fund of the 
Chinese Academy of Science under the authority of the Ministry of Finance 
(STZ-00-19). 

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Amphipoda: Gammaridae). Acta Zootaxonomica Sinica 28 (4): 621-628. 
Morino, H. & Whitman, N. 1995. A new species of the genus Gammarus (Crustacea: Amphi- 
poda) from fresh waters of Russian Far East. Publications of Itako Hydrobiological 

Station 8: 27-36. 



284 Z. E. HOU ET AL. 



Schaferna, K. 1923. Amphipoda balcanica, spolu s poznâmkami o jinych sladkovodnich 
Amphipodech. Vestnik Kralovske Ceske Spolecnosti Nauk, 1921-1922 (12): 1-111. 

Stock, J. H., Mirzajani, A. R., Vonk, R., Naderi, S. & Kiabi, B. H. 1998. Limnic and brackish 
water Amphipoda (Crustacea) from Iran. Beaufortia 48 (9): 173-234. 

Tattersall, W. M. 1924. Zoological results of the Percy Sladen Trust expedition to Yunnan, 
under the leadership of professor J. W. Gregory, F. R. S. (1922). Journal and Pro- 
ceedings of the Asiatic Society of Bengal (New series) 19 (9): 429-435. 



Revue suisse de Zoologie 111 (2): 285-301; juin 2004 



Neue orientalische Arten der Gattung S te nus Latreille 
aus dem Genfer Museum (Coleoptera: Staphylinidae) 
284. Beitrag zur Kenntnis der Steninen 

Volker PUTHZ 

c/o Limnologische Fluß-Station, MPI-Limnologie, Damenweg 1, D-36110 Schlitz, 

Deutschland. E-Mail: Stenus.Puthz@t-online.de 



New Oriental species of the genus S tenus Latreille from the Geneva 
Museum (Coleoptera: Staphylinidae). 284 th Contribution to the know- 
ledge of Steninae. - Description of 11 new species: Stenus (Hemistenus) 
agostii sp. n. (Sumatra), S. (Hemistenus) croceipennis sp. n. (Sumatra), S. 
(s. str.) cuccodoroi sp. n. (Papua New Guinea), S. (Hypostenus) exsecratus 
sp. n. (Sulawesi), S. (Hypostenus) kaibesarensis sp. n. (Moluccas: Kai 
Besar), S. (Hypostenus) kerincimontis sp. n. (Sumatra), S. (Hemistenus) kur- 
batovi sp. n. (Burma), S. (Hypostenus) radulipenis sp. n. (Thailand), S. (s. 
str.) sannifer (Java, Lombok), S. (Hemistenus) schwendingeri sp. n. 
(Thailand) and S. (Hypostenus) tanimbarensis sp. n. (Moluccas: Tanimbar). 

Key- words: Coleoptera - Staphylinidae - Stenus - taxonomy. 



EINLEITUNG 

Im umfangreichen Staphylinidenmaterial des Genfer Naturhistorischen 
Museums fand sich eine Anzahl neuer Stenus- Arten aus der Orientalis. Von diesen 
werden hier 11 beschrieben. Sie gehören zu verschiedenen Verwandtschaftsgruppen, 
wie jeweils in den Einzelbeschreibungen angegeben. Bemerkenswert ist die Tatsache, 
dass die beiden neuen von den Molukken beschriebenen Arten zeigen, dass die Gruppe 
des Stenus gigas L. Benick östlich der Weber-Linie (vgl. Joly, 2003) eine reiche 
Differenzierung erfahren hat. - Sofern nicht anders angegeben, befinden sich Holo- 
typen und Paratypen im Muséum d'histoire naturelle de Genève und Paratypen in 
meiner Sammlung. 

Wie in meinen anderen Arbeiten gelten auch hier die folgenden Abkürzungen: 
aE = average distance between eyes, mittlerer Augenabstand; HT = Holotypus; 1E1 = 
greatest length of elytra, größte Elytrenlänge; IE = length of eyes, Augenlänge; IP = 
length of pronotum, Pronotumlänge; IS = length of suture, Nahtlänge; PM = propor- 
tional measurements, Proportionsmaße (1 Einheit = 0,0252 mm); PT = Paratypus; wEl 
= greatest width of elytra, größte Elytrenbreite; wH = width of head, Kopfbreite; wP = 
width of pronotum, Halsschildbreite. 



Manuskript angenommen am 17.10.2003 



286 v. PUTHZ 

SYSTEMATIK 

Stenus (s. str.) sannifer sp. n. 

Material: o*-Holotypus und 2 5 9-Paratypen: [Indonesien]: Java: Maribaya, 22 km NE 
Bandung, 27.VII.1984, J. Robert (MHNG, 1 PT coll. Puthz); 1 3 -Paratypus: [Indonesien]: 
Lombok: Senaro, N slope of Rinjani, 1100 m, 2.- 5. II. 1984, Bolm (Staatliches Museum für 
Naturkunde, Stuttgart) ; 4 9 9-Paratypen: Mt. Rinjani, Sebnaro, 400 m, waterfalls, 5.XI.1991, 
Lobi (MHNG, 1 PT coll. Puthz). 

Beschreibung: Länge 2,5-3,1 mm (Vorderkörperlänge: 1.6 mm). Schwarz, 
mäßig glänzend mit leichtem Messingschimmer, Vorderkörper sehr grob und gedrängt- 
dicht bis rugos punktiert, Abdomen grob und sehr dicht punktiert; Beborstung kurz, 
wenig auffällig. Fühler bräunlichgelb, Kiefertaster gelb, Beine bräunlichgelb, die 
Schenkelspitzen angedunkelt. Clypeus schwarz, Oberlippe dunkelbraun, ziemlich 
dicht beborstet. 

PM des HT und des c?-PT von Lombok (in Klammern): wH: 27,5 (30); aE: 16 
(17); wP: 21 (23); IP: 20 (23); wEl: 27,5 (31,5); 1E1: 25 (28); IS: 19 (22). 

Männchen: Schienen ohne Dornen, Hinterschienen in ihren apikalen zwei 
Dritteln leicht gebogen und innen abgeflacht. Vordersternite ohne Auszeichnungen. 7. 
Sternit vor dem Hinterrand feiner und dichter als an den Seiten punktiert und beborstet. 
8. Sternit (Fig. 2). 9. Sternit apikolateral mit spitzem Zahn. 10. Tergit breit abgerundet. 
Aedoeagus (Fig. 1), der breite und vergleichsweise kurze Medianlobus ist vorn 
nach einer gewellten Kontur in eine kurze, dreieckige Spitze verengt; im Innern 
werden zwei kräftige, distal gesägte, median verbundene Ausstülphaken und eine 
dünne Tube deutlich. Die Parameren sind viel länger als der Medianlobus und tragen 
mehrere Gruppen ziemlich langer Borsten. 

Weibchen: 8. Sternit zur Hinterrandmitte leicht vorgezogen, abgerundet. 
Valvifer apikolateral mit spitzem Zahn. 10. Tergit breit abgerundet. Die Spermatheka 
besteht aus einem langen, dünnen, dreimal gewundenen Schlauch und ist insgesamt 
fast so lang wie ein Valvifer. 

Bemerkungen: Diese neue Art gehört in die Gruppe des S. javanus Bernhauer 
(Puthz, 1980: 29) und ist hier die Schwesterart des S. javanus. Äußerlich kann ich sie 
nicht sicher von jenem trennen, zumal sie auch skulpturell variabel ist: die oben 
genannten Proportionsmaße des HT und des â von Lombok (beide haben identische 
Aedoeagen) zeigen schon, welche Unterschiede hier auftreten können. Beim HT 
handelt es sich um ein schlankeres, deutlich glänzendes und am Vörderkörper kaum 
rugos punktiertes Exemplar, die Stücke aus Lombok dagegen sind robuster, weniger 
glänzend und am Vörderkörper erheblich rugoser skulptiert. Zu S. javanus (dessen HT 
ich ebenso wie den des synonymen S. brachycephalus Cameron noch einmal unter- 
sucht habe) kann ich keine sicheren ektoskelettalen Unterscheidungsmerkmale 
angeben. Der Aedoeagus beider Arten zeigt aber ihren Unterschied (vgl. Fig. 4). Von 
S. sannator Puthz (Nord-Indien, Nepal), dessen Aedoeagus einen ähnlichen Umriss 
besitzt, trennt man die neue Art durch weniger tief ausgerandetes 8. Sternit des 
Männchens (vgl. Fig. 3) und durch den anders gestalteten Ausstülpapparat des Median- 
lobus (vgl. Abb. 6, Puthz, 1991), von S. kamhaengi Rougemont durch fehlende 
Elytrenmakeln, von S. spongifera Cameron, S. riukiuensis Puthz und S. sannio Puthz 
durch weniger breite Stirn und durch seinen vorn stumpfwinklig (also nicht gerundet-) 
verengten Medianlobus. 



NEUE ORIENTALISCHE STENUS ARTEN 287 

Etymologie: Ich nenne die neue Art sannifer = grimassentragend, einmal um 
ihre Verwandtschaft mit den ähnlich benannten Spezies zu signalisieren, dann aber 
auch um auszudrücken, dass sie äußerlich merklich variabel ist. 

Stenus (s. str.) cuccodoroi sp. n. 

Material: c?-Holotypus und 3 6 6-, 2 9 Ç-Paratypen: Papua Neuguinea: Morobe, Biaru 
Rd., Mt. Saredomo, 2450 n, 9.VI.1992, G. Cuccodoro # 20A (MHNG, 2 PTT coli. Puthz). 

Beschreibung: Länge: 2,7-3,7 mm (Vorderkörperlänge: 1,6-1,8 mm). Preußisch- 
blau-metallisch, glänzend, Vorderkörper sehr grob und sehr dicht punktiert, Abdomen 
ziemlich grob und mäßig dicht punktiert; Beborstung kurz, wenig auffällig. Fühler 
hellbraun, die Keule dunkler, Kiefertaster gelb, das 3. Glied wenig dunkler, Beine 
kastanienbraun, die Schenkelspitzen etwas dunkler. Clypeus metallisch bis schwarz, 
Oberlippe schwarzbraun, beide ziemlich dicht beborstet. 

PM des HT und eines cT-PT in Klammern: wH: 27,5 (27,8); aE:16 (16); wP: 
21,8 (23), IP: 23 (25); wEl: 30 (35); 1E1: 27 (33,5); IS: 22 (28). 

Männchen: Beine ohne Auszeichnungen. Metasternum abgeflacht, sehr grob, 
dicht punktiert, Hinterhüftumrandung glatt. Vordersternite ohne Auszeichnungen. 7. 
Sternit in der hinteren Mitte sehr fein und sehr dicht punktiert und beborstet, Hinter- 
rand sehr flach ausgerandet. 8. Sternit mit breitrunder Ausrandung etwa im hinteren 
Zwölftel. 9. Sternit apikolateral mit spitzem Zahn. 10. Tergit breit abgerundet. 
Aedoeagus (Fig. 5), Medianlobus stumpf-dreieckig verengt und im Innern mit 
dünntubigem Innensack sowie starken Ausstülphaken. Parameren viel länger als der 
Medianlobus, an ihren Spitzen nur mit zwei subapikalen Borsten. 

Weibchen: 8. Sternit breit abgerundet. Valvifer apikolateral mit spitzem Zahn. 
10. Tergit abgerundet. 

Bemerkungen: Diese neue Art gehört in die Gruppe des S. toxopei Cameron 
(Puthz, 1971: 448f.) und ist hier die Schwesterart des S. delicatus Puthz, dem sie in fast 
allen Punkten grundsätzlich ähnelt. Sie unterscheidet sich jedoch von ihm durch 
schmäleren Kopf (deutlich schmäler als die Elytren) mit flacherer, weniger tief einge- 
senkter Stirn mit etwas gröberer Punktierung (die Punkte erreichen gut den apikalen 
Querschnitt des 2. Fühlergliedes), durch gröbere Pronotumpunktierung (die Pronotum- 
punktierung ist nur wenig feiner als diejenige der Elytren, bei S. delicatus ist der 
Unterschied viel auffälliger) sowie durch den Innenbau (Hakenumriss) des Median- 
lobus. Von S. delicatulus Puthz unterscheidet sich die neue Art durch schmäleren Kopf, 
breitere Elytren, weniger grob punktiertes Abdomen und den Aedoeagus. In ihren 
Proportionen sind die Stücke der Typenserie merklich variabel: ein c?-Paratypus fällt 
durch erheblichere Größe und breitere Elytren auf (vgl. o.). 

Etymologie: Ich dediziere diese schöne neue Art ihrem Sammler, Herrn Dr G. 
Cuccodoro (Genf). 

Stenus (Hypostenus) kerincimontis sp. n. 

Material: d-Holotypus und 1 9-Paratypus: [Indonesien]: Sumatra: Jambi, Mt. 
Kerinci, 3300 m, 12.XI.1989, Agosti, Lobi & Burckhardt # 12a (MHNG, PT coli. Puthz): 1 ?- 
Paratypus: ibidem 2100 m, 14.XI.1989, idem # 16 (MHNG). 

Beschreibung: Länge: 2,8-3,6 mm (Vorderkörperlänge: 1,7 mm). Brachypter, 
schwarz, Elytren mit bräunlichem Anflug, schwach glänzend, Stirn ziemlich grob und 



288 



V. PUTHZ 








Fig. 1-5 
Ventralansicht der Aedoeagen (1, 4, 5) und 8. Sternit der Männchen (2, 3) von Stenus (s. str.) san- 
nifer sp. n. (PT: 1, 2), S. sannator Puthz (3), S.javanus Bernhauer (HT von S. brachycephalus 
Cameron, 4) und S. (s. str.) cuccodoroi sp. n. (FT, 5).- Maßstab = 0,1 mm. 



dicht punktiert, Pronotum grob und dicht punktiert, Elytren sehr grob und dicht punk- 
tiert, Abdomen ziemlich grob (vorn) bis fein (hinten), überall dicht punktiert. Fühler 
gelb, die Keule gebräunt, Kiefertaster gelb bis rötlichgelb, Beine bräunlichgelb, 
Schenkel- und Tarsengliedspitzen kaum dunkler. Clypeus schwarz, Oberlippe 
schwarzbraun, heller gesäumt, beide spärlich beborstet. Paraglossen oval. 

PM des HT: wH: 35; aE: 17; wP: 25; IP: 26; wEl: 31; LEI: 29; IS: 20. 

Männchen: Beine ohne Auszeichnungen. Metasternum leicht eingedrückt, grob 
und dicht auf eng genetztem Grund punktiert, Hinterhüftumrandung innen mit 
mehreren mäßig feinen Punkten. Vordersternite einfach. 7. Sternit in der hinteren Mitte 
sehr fein und dicht punktiert und beborstet, Hinterrand kaum erkennbar flach aus- 
gerandet. 8. Sternit mit kleinem rundem Ausschnitt etwa im hinteren Achtzehntel. 9. 
Sternit apikolateral mit spitzem Zahn. 10. Tergit breit abgerundet. Aedoeagus 
(Fig. 8), Medianlobus vorn konvex verengt, im Innern mit stark skierotisiertem 
Innensack und median kurz verbundenen Ausstülphaken (Fig. 10). Parameren viel 
länger als der Medianlobus, in ihrer Spitzenhälfte lang verschmälert, mit zwei 
Borstengruppen (7, 5). 

Weibchen: 8. Sternit zur Hinterrandmitte dreieckig vorgezogen. Valvifer apiko- 
lateral mit spitzem Zahn. 10. Tergit breit abgerundet. Spermatheka (Fig. 9), etwas kürz- 
er als der stärker skierotisierte distale Teil des Valvifers. 



NEUE ORIENTALISCHE STENUS ARTEN 



289 




Fig. 6-13 
Ventralansicht der Aedoeagen, zum Teil ohne Innenkörper, weil ausgestülpt (6, 8, 13), 9. Sternit 
des Männchens (7, 12), Spermatheka (9) und Ausstülphakenapparat des Medianlobus (10, 11) 
von Stenus (Hypostenus) exsecratus sp. n. (HT, 6, 7), S. (Hypostenus) kerincimontis sp. n. 
(8-10), S. (Hemistenus) schwendingeri sp. n. (HT, 11-13).- Maßstab = 0,1 mm (6= 7-10; 13= 11, 
12). 



Kopf deutlich breiter als die Elytren, Stirn ziemlich schmal mit deutlichen 
Längsfurchen, Mittelteil gut so breit wie jedes der Seitenstücke, breitrund erhoben, 
erreicht fast das Augeninnenrandniveau. Punktierung ziemlich grob und sehr dicht, 
mittlerer Punktdurchmesser gut so groß wie der mittlere Querschnitt des 3. Fühler- 
gliedes, Punktabstände fast überall kleiner als die Punktradien. Fühler schlank, zurück- 
gelegt knapp den Pronotumhinterrand überragend, vorletzte Glieder etwa eineinhalb 
mal so lang wie breit. Pronotum kaum länger als breit, seitlich kräftig gewölbt, hinten 
deutlich konkav verengt. Eine abgekürzte mittlere Längsfurche und seitliche Schräg- 
eindrücke mäßig deutlich. Punktierung gröber als auf der Stirn, dicht, mittlerer Punkt- 
durchmesser gut so groß wie der größte Querschnitt des 3. Fühlergliedes, aber kleiner 
als der apikale Querschnitt des 2. Fühlergliedes, die glänzenden Punktzwischenräume 
fast überall (wenig) kleiner als die Punktradien. Elytren rechteckig mit ausgeprägten 
Schultern, deutlich schmäler als der Kopf und deutlich breiter als lang, die Seiten 
etwas, gerade, erweitert, Hinterrand tiefrund ausgerandet; keine Eindrücke. Punk- 
tierung sehr grob und dicht, aber getrennt, mittlerer Punktdurchmesser so groß wie der 
mittlere Querschnitt des 2. Fühlergliedes, die genetzten Punktzwischenräume kleiner 
als die Punktradien. Abdomen stark gewölbt, ellipsoid, die Segmente ohne deutliche 
seitliche Trennungslinie, wenn auch die unterschiedliche Anordnung der dorsalen und 
der ventralen Punktierung eine solche vortäuscht. Basale Querfurchen der ersten 



290 V. PUTHZ 

Segmente ziemlich tief, 7. Tergit mit rudimentärem apikalem Hautsaum (ungeflügelte 
Art). Die Punktierung der ersten Tergite ist jeweils vorn etwa so grob wie auf der Stirn, 
dicht, nahe dem Hinterrand feiner, das 7. Tergit ist fein und ziemlich dicht punktiert, 
die Punkte sind etwas kleiner als eine Augeninnenrandfacette, ihre Abstände gut punk- 
tgroß. Das 10. Tergit ist fein und mäßig dicht punktiert. Beine schlank, Hintertarsen 
mehr als zwei Drittel schienenlang, ihr 1. Glied ist deutlich länger als die beiden fol- 
genden zusammengenommen, erheblich länger als das Klauenglied; das 4. Glied ist 
lang und schmal gelappt. Die Stirn zeigt allenfalls Netzung s spuren, das Pronotum ist 
ungenetzt, Elytren und Abndomen mehr oder weniger deutlich genetzt. 

Bemerkungen: Diese neue Art ist genitaliter den Spezies S. voraginosus L. 
Benick (Java) und S. exsecratus sp. n. (Sulawesi) sehr ähnlich, vom ersteren unter- 
scheidet sie sich durch robusteren Bau und die weniger grobe Punktierung des Vorder- 
körpers, vom zweiten durch kürzere Elytren, gröbere Punktierung von Pronotum und 
Elytren und weniger dicht punktiertes Abdomen, von beiden durch die Sexual- 
charaktere. Von allen anderen orientalischen Hypostenen durch die Kombination der 
Merkmale: einfaches 10. Tergit, apikolateral spitzes 9. Sternum, fehlende Elytren- 
makeln, breiten Kopf, helle Beine und ihre geringe Größe. Sie ähnelt habituell den 
(Hemistenus-) Arten S. aeneopullus Puthz, S. iniustus Puthz und S. vulcanus Rouge- 
mont, die aber alle nur eine flache Stirnmitte (keinen breit erhobenen Mittelteil) 
besitzen; von S. despectus L. Benick unterscheidet sie sich durch nahezu rechteckige, 
also nicht lang-trapezoide Elytren, von allen diesen durch apikolateral einspitziges 9. 
Sternum und die fehlende Seitenrandung des Abdomens, schließlich von S. pilicornis 
Fauvel, der ebenfalls auf dem Mt. Kerinci lebt, durch feinere Stirnpunktierung, 
ungerandetes Abdomen und die Sexualcharaktere. 

Etymologie: Der Name bezeichnet die Herkunft der neuen Art. 

Stenus (Hypostenus) exsecratus sp. n. 

Material: <?-Holotypus: [Indonesien]: Sulawesi: Gunang Mogogonia Pa 1080 m, 
Utara, X.1985, R. W. Homabrook (coli. Puthz im MHNG). 

Beschreibung: Länge: 2,6-3,4 mm (Vorderkörperlänge: 1,6 mm). Schwarz mit 
leichtem Brauneinschlag, schwach schimmernd, grob und überall sehr dicht punktiert, 
Beborstung kurz, nur am Abdomen deutlich. Fühler gelb, die Keule wenig dunkler, 
Kiefertaster gelb, Beine gelblich, die Schenkel zur Spitze gelblichbraun. Clypeus 
schwarz, Oberlippe dunkelbraun, heller gesäumt, beide spärlich beborstet. 

PM des HT: wH: 29; aE: 15; wP: 22; IP: 22; wEl: 29; 1E1: 29; IS: 23. 

Männchen: Beine ohne Auszeichnungen. Metasternum grob und dicht auf 
glänzendem Grund punktiert. Vordersternite einfach. 7. Sternit median abgeflacht und 
daselbst sehr fein und dicht punktiert und beborstet. 8. Sternit am Hinterrand kaum 
erkennbar flach ausgerandet. 9. Sternit (Fig. 7). 10. Tergit breit abgerundet. 
Aedoeagus (Fig. 6), Medianlobus vorn abgerundet, im Innern mit stark skiero- 
tisiertem Innensack und großen, median kurz verbundenen Ausstülphaken,. Parameren 
viel länger als der Medianlobus, zur Spitze lang verschmälert und mit zwei getrennten 
Borstengruppen. 

Weibchen: unbekannt. 



NEUE ORIENTALISCHE STENUS ARTEN 291 

Kopf etwa so breit wie die Elytren, Stirn schmal mit deutlichen Längsfurchen, 
Mittelteil so breit wie jedes der Seitenstücke, deutlich, aber wenig, breitrund erhoben, 
erreicht nicht die Höhe der Augeninnenränder. Punktierung grob und überall sehr dicht, 
mittlerer Punktdurchmesser gut so groß wie der mittlere Querschnitt des 3. Fühler- 
gliedes, die scharfen Punktzwischenräume viel kleiner als die Punktradien. Fühler 
schlank, zurückgelegt den Hinterrand des Pronotums erreichend, vorletzte Glieder 
wenig länger als breit. Pronotum so lang wie breit, vorn seitlich kräftig konvex, hinten 
deutlich eingeschnürt, mit einer vorn und hinten abgekürzten deutlichen Mittelfurche, 
seitlich kaum uneben. Punktierung so grob und ebenso dicht wie auf der Stirn, die 
Punkte sind aber tiefer eingestochen. Elytren so breit wie der Kopf, quadratisch, so 
lang wie breit, Schultern rechteckig, Schultereindrücke und ein vorderer Nahteindruck 
deutlich. Punktierung deutlich etwas gröber als am Pronotum, überall sehr dicht, 
mittlerer Punktdurchmesser so groß wie der apikale Querschnitt des 2. Fühlergliedes. 
Abdomen breit-zylindrisch, eine Seitenrandung nur skulpturell angedeutet, nicht klar 
ausgeprägt, basale Quereinschnürungen der ersten Segmente tief, 7. Tergit mit 
deutlichem apikalem Hautsaum (die Art ist makropter). Punktierung vorn etwa so grob 
wie auf der Stirn, nach hinten deutlich feiner, überall sehr dicht, auf dem 7. Tergit sind 
die Punkte wenig feiner als eine Augeninnenrandfacette, ihre Abstände fast so groß wie 
die Punkte, das 10. Tergit ist sehr fein und spärlich punktiert. Beine schlank, Hinter- 
tarsen drei Viertel schienenlang, ihr 1 . Glied fast so lang wie die drei folgenden zusam- 
mengenommen, erheblich länger als das Klauenglied; 4. Glied schmal und tief gelappt. 
Stirn mit Netzungsspuren, Pronotum und Elytren zwischen den Punkten glatt, das 
gesamte Abdomen deutlich genetzt. 

Bemerkungen: Diese neue Art ist (genitaliter) die Schwesterart des S. voragi- 
nosus L. Benick. Sie unterscheidet sich von ihm äußerlich sofort durch ihre größeren 
Elytren und die erheblich feinere und vor allem am Abdomen viel dichtere Punktierung 
sowie durch das seitlich ungerandete Abdomen (S. voraginosus besitzt eine sehr dünne, 
aber vollständige Seitenlinie, wenn auch keine Trennung in Tergite und Sternite). Von 
allen orientalischen {Hypostenus-) Arten unterscheidet sie sich ebenso wie S. kerinci- 
montis (vgl. o.), von diesem durch größere Elytren, viel dichtere Abdominal- 
punktierung und die Sexualcharaktere. Habituell ähnelt sie den {Hemistenus-) Spezies 
S. absurdus Puthz, S. guru Puthz, S. inconspicuus Cameron und S. perfidiosus Puthz, 
die auch alle ein apikolateral spitzes 9. Sternit besitzen, die aber entweder eine gröber 
punktierte oder flache Stirn und/oder gröber punktierte Abdomina aufweisen, ganz 
abgesehen von den Sexualcharakteren. 

Etymologie: Ich nenne diese Art „verwünscht" (lat. exsecratus), weil sie sich 
äußerlich so stark von ihren Nahverwandten unterscheidet und man ihre Verwandt- 
schaft erst nach Genitaluntersuchung überrascht erkennt. 

Stenus (Hypostenus) radulipenis sp. n. 

Material: c?-Holotypus: THAILAND: Taksin Maharat National Park [westlich Tak], 
1000 m, 9.II.1993, Schwendinger. 

Beschreibung: Länge: 4,0-5,0 mm (Vorderkörperlänge: 2,4 mm). Schwarz, 
Elytren mit großer, ovaler, orangegelber Makel, Vorderkörper fast matt, Abdomen 
etwas glänzend, Stirn grob und sehr dicht punktiert (keine Glättungen), Pronotum sehr 



292 



V. PUTHZ 






Ventralansicht der Aedoeagen von Stenus (Hypostenus) kaibesarensis sp. n. (HT, 14), S. 
(Hypostenus) tanimbarensis sp. n. (HT, 15) und S. (Hemistenus) kurbatovi sp. n. (HT, 16).- 
Maßstab = 0,1 mm. 



grob, sehr tief eingestochen und sehr dicht, überall kurz-zusammenfließend skulptiert, 
außerdem mit mehreren tiefen Eindrücken, Elytren sehr grob und äußerst dicht, leicht 
zusammenfließend punktiert, Abdomen vorn grob und sehr dicht, hinten wenig fein 
und dicht auf netzungsfreiem Grund punktiert; Beborstung kurz, wenig auffällig. 
Fühlerbasis rötlichgelb, die Keule dunkler, Kiefertaster rötlichgelb, Beine rötlichgelb, 
die Schenkelspitzen etwas dunkler. Clypeus schwarz, Oberlippe dunkelbraun bis 
(vorn) heller, Clypeus dicht, Oberlippe dünner beborstet. 

PM des HT: wH: 47; aE: 23; wP: 33; IP: 34; wEl: 46; LEI: 44; IS: 32. 

Männchen: Mittelschienen mit kleinem Apikaldorn, Hinterschienen mit 
kleinem Präapikaldorn. Vordersternite ohne Auszeichnungen, 7. Sternit median feiner 
und dichter als an den Seiten punktiert und beborstet, Hinterrand kaum erkennbar flach 
ausgerandet. 8. Sternit mit geschwungen-spitzwinkligem Ausschnitt fast im hinteren 
Fünftel. 9. Sternit apikolateral mit kurzem, spitzem Zahn. 10. Tergit sehr breit ab- 
gerundet. Aedoeagus (Fig. 17), die Apikaipartie des Medianlobus etwa 
lanzettlich, leicht dorsad gebogen und dicht mit kurzen Sinnesborsten versehen. Im 
Innern wird die für die guttalis-Gruppe kennzeichnende dreieckige Spange erkennbar. 
Parameren deutlich länger als der Medianlobus, zur Spitze erweitert und daselbst mit 
zahlreichen mäßig langen Borsten versehen. 

Weibchen: unbekannt. 

Bemerkungen: Diese neue Art gehört in die guttalis-Gmppe (Puthz, 1988: 635) 
und ist hier die Schwesterart des S. subguttalis Puthz. In ihren ektoskelettalen Merk- 
malen ähnelt sie mehreren Arten ihrer Gruppe stark, so dass es zu ihrer Charak- 



NEUE ORIENTALISCHE STENUS ARTEN 



293 




Fig. 17-19 
Ventralansichten der Aedoeagen von Stenus (Hypostenus) radulipenis sp. n. (HT, 17), S. 
(Hemistenus) croceipennis sp. n. (PT, 18) und S. (Hemistenus) agostii sp. n. (PT, 19).- Maßstab 
= 0,1 mm. 



terisierung ausreicht, neben einer Kurzbeschreibung die wesentlichen Unter- 
scheidungsmerkmale zu nennen. Sie unterscheidet sich von ihren (Hypostenus-) Ver- 
wandten S. asperrimus L. Benick, S. bivulneratus Motschulsky, S. burckhardti 
Rougemont, S. guttalis Fauvel und S. iniquus L. Benick durch ungenetztes Abdomen, 
von S. asperrimus, S. iniquus und S. subiniquus Puthz durch ihre Elytrenmakeln, 
schließlich von S. subguttalis Puthz und S. variipennis Rougemont durch breitere und 
längere, mehr rundlich-rechteckige (also nicht trapezoide) Elytren, länglichere (ovale) 
Elytrenmakeln und weniger grob und dicht punktierte Abdomenspitze (Punktabstände 
bei der neuen Art meist etwa so groß wie die Punkte), von allen genannten Spezies 
durch den Aedoeagus. 

Etymologie: Weil die Apikaipartie des Medianlobus an eine Radula erinnert, 
nenne ich die neue Art radulipenis. 



Stenus (Hypostenus) kaibesarensis sp. n. 

Material: <?-Holotypus: [Indonesien]: Maluc: Kai Besar G., Tukrau, 300 m, leaf litter. 
7.IX.1991, D. Agosti., F911074, # 8 (MHNG); 1 c?-Paratypus: Kai Besar: Bombay (E. of Elat). 
G. Dab, 3.IX.1991, D. Agosti, F911018, # 4 (coll. Puthz). 



294 v. PUTHZ 

Beschreibung: Länge: 4,5-5,7 mm (Vorderkörperlänge: 2,3 mm). Brachypter, 
blaumetallisch, ziemlich glänzend, Stirn grob und ziemlich weitläufig punktiert, 
Pronotum und Elytren sehr grob und sehr dicht, aber getrennt punktiert, Abdomen grob 
und ziemlich dicht punktiert; Beborstung kurz, wenig auffällig. Fühler hellbraun, die 
Keule wenig dunkler, Kiefertaster rötlichgelb, Beine rötlichgelb, die Knie und die 
Tarsen wenig dunkler. Clypeus blaumetallisch bis schwarzbraun, Oberlippe dunkel- 
braun, heller gesäumt, beide mäßig dicht beborstet. 

PM des HT: wH: 45,5; aE: 26; wP: 31,5; IP: 35; wEl: 41; 1E1: 34,5; IS: 22. 

Männchen: Beine ohne Auszeichnungen. Metasternum abgeflacht, grob und 
ziemlich dicht punktiert. Vordersternite grob und dicht punktiert. 7. Sternit in der hin- 
teren Mitte feiner (aber immer noch ziemlich grob) und dichter als an den Seiten punk- 
tiert. 8. Sternit mit geschwungen-stumpfwinkligem Ausschnitt etwa im hinteren Elftel. 
9. Sternit apikolateral mit spitzem Zahn. 10. Tergit abgerundet. Aedoeagus (Fig. 
14) mit spitzwinklig verengtem Medianlobus, dieser vorn mit einem ventralen Feld 
dicht stehender Sinnesborsten. Parameren kürzer als der Medianlobus, apikal etwas 
verbreitert und daselbst mit ca. 10 kräftigen Borsten. 

Weibchen: unbekannt. 

Kopf deutlich breiter als die Elytren, Stirn mäßig breit mit flachen Längsfurchen 
und einem kaum erhobenen Mittelteil, der etwas schmäler als jedes der Seitenstücke 
ist. Punktierung grob, mittlerer Punktdurchmesser gut so groß wie der basale 
Querschnitt des 3. Fühlergliedes, auf den Seiten meist dicht (Punktabstände hier größer 
als die Punktradien), neben dem hinteren Augeninnenrand mit kleiner punktfreier 
Partie, Stirnmitte nur mit wenigen, zerstreuten Punkten, überwiegend punktfrei. Fühler 
kurz, zurückgelegt bis ins hintere Pronotumdrittel reichend, vorletzte Glieder einein- 
halb mal so lang wie breit. Pronotum breit tonnenförmig, deutlich etwas länger als 
breit, seitlich wenig gewölbt, hinten nur flach konkav verengt, ohne Eindrücke. 
Punktierung sehr grob und sehr dicht, der mittlere Punktdurchmesser ist nur wenig 
kleiner als der größte Querschnitt des 2. Fühlergliedes, die Punktzwischenräume sind 
bis auf wenige Ausnahmen überall viel schmäler als die Punktradien. Elytren 
trapezoid, viel schmäler als der Kopf, erheblich breiter als lang, Schultern schräg, 
Seiten kräftig nach hinten erweitert, Hinterrand tiefrund ausgerandet; ein flacher 
Schultereindruck erkennbar. Punktierung ähnlich grob und dicht wie am Pronotum. 
Abdomen stark gewölbt, fast zylindrisch, die Segmente 4-6 zeigen nur eine ange- 
deutete seitliche Trennungslinie, die basalen Quereinschnürungen der ersten Segmente 
sind sehr tief, das 7. Tergit trägt einen reduzierten apikalen Hautsaum. Die Punktierung 
ist grob und ziemlich dicht, vorn nur wenig gröber als hinten, auf dem 3. Tergit sind 
die Punkte so groß wie der basale Querschnitt des 3. Fühlergliedes, ihre Abstände etwa 
punktgroß, auf dem 7. Tergit sind die etwas längs-ausgezogenen Punkte kaum kleiner, 
ihre Abstände gut punktgroß, das 10. Tergit ist wenig fein und wenig dicht punktiert. 
Beine schlank, die Hintertarsen nicht ganz drei Fünftel schienenlang, ihr 1. Glied ist et- 
was kürzer als die drei folgenden zusammengenommen, deutlich länger als das 
Klauenglied; das 4. Glied ist tief gelappt. Die Stirn und der überwiegende Teil des 
Abdomens sind deutlich genetzt, Pronotum und Elytren sowie das 8. Tergit zeigen nur 
Netzungsspuren, 10. Tergit ungenetzt. 

Bemerkungen: Diese neue Art gehört in die Gruppe des S. gigas L. Benick 
(Puthz, 1971: 449) und hier in die Nähe des S. magnificus L. Benick. Sie unterschiedet 



NEUE ORIENTALISCHE STENUS ARTEN 295 

sich von all ihren Nahverwandten durch ihre kurzen, breiten Elytren, ihre sehr grobe 
Vorderkörperpunktierung, ihre Netzung und durch den Aedoeagus. 

Etymologie: Der Name dieser Art bezeichnet ihre Herkunft von der 
Molukkeninsel Kai Besar. 

Stenus (Hypostenus) tanimbarensis sp. n. 

Material: <?-Holotypus und ?-Paratypus: [Indonesien]: Maluc: Tanimbar Island: 
Yamdena Domaki, NW of Saumlaki, 18.IX.1991, D. Agosti, F911146. 

Beschreibung: Länge: 5,0-6,2 mm (Vorderkörperlänge: 2,5 mm). Brachypter, 
blaugün bis grasgrünmetallisch, stark glänzend, Stirn mäßig fein und ziemlich weitläu- 
fig punktiert, Pronotum und Elytren grob und dicht punktiert, Abdomen mäßig grob 
und ziemlich weitläufig punktiert; Beborstung kurz, wenig auffällig. Fühler hellbraun, 
die Keule etwas dunkler, Kiefertaster rötlichgelb, Beine bräunlichgelb, Schenkel- 
spitzen und Tarsen wenig dunkler. Clypeus metallisch bis schwarz, Oberlippe dunkel- 
braun bis heller, beide mäßig dicht beborstet. 

PM des HT: wH: 48; aE: 24; wP: 33; IP: 35; wEl: 42; 1E1: 38; IS: 24. 

Männchen: Beine ohne Auszeichnungen. Metasternum abgeflacht, grob und 
mäßig dicht punktiert. 6. Sternit median feiner und dichter als an den Seiten punktiert 
und beborstet. 7. Sternit median sehr fein und sehr dicht punktiert und beborstet. 8. 
Sternit mit stumpfwinkligem Ausschnitt etwa im hinteren Achtel. 9. Sternit apiko- 
lateral mit spitzem Zahn. 10. Tergit zum Hinterrand rundlich vorgezogen. 
Aedoeagus (Fig. 15) ähnlich dem des S. kaibesarensis, Medianlobus aber etwas 
schlanker, apikoventral mit zwei Sinnesborstenfeldern, Parameren stärker und länger 
beborstet. 

Weibchen: 8. Sternit schmal abgerundet. Valvifer apikolateral mit spitzem 
Zahn. 10. Tergit breit abgerundet. 

Kopf ähnlich wie bei S. kaibesarensis, Stirn jedoch schmäler, Mittelteil eben- 
falls nur sehr flach gewölbt, Punktierung aber viel feiner: die größten Punkte sind 
deutlich kleiner als der basale Querschnitt des 3. Fühlergliedes, etwa so groß wie eine 
Augenfacette, die kleinsten (auf der hinteren Partie der Seitenstücke) erheblich kleiner; 
die Stirnmitte ist überwiegend punktfrei, auf den Seitenstücken stehen die Punkte im 
mittleren Bereich dicht, ihre Abstände werden aber nach vorn und vor allem nach 
hinten erheblich größer. Fühler und Pronotumumriss ähnlich wie bei der verglichenen 
Art, die Punktierung des Pronotums zwar grob, aber viel feiner als bei S. kaibesaren- 
sis, der mittlere Punktdurchmesser liegt etwas über dem basalen Querschnitt des 3. 
Fühlergliedes, die glänzenden Punktzwischenräume sind oft kaum schmäler als die 
Punktradien, ausnahmsweise auch einmal punktgroß. Die Elytren sind ebenfalls 
trapezoid und im Umriss ähnlich wie bei S. kaibesarensis, ihre Punktierung aber 
weniger grob und nicht so gedrängt, die Punkte leicht quer-ausgezogen, der mittlere 
Punktdurchmesser entspricht etwa dem größten Querschnitt des 3. Fühlergliedes, die 
Punktabstände sind auf der Scheibe deutlich etwas schmäler als die Punktradien, 
seitlich so groß oder auch größer; in der Elytrenhinterhälfte stehen die Punkte in 
leichten Querreihen angeordnet. Abdomen prinzipiell wie bei S. kaibesarensis, auch 
die Punktierung. Auch die Beine etwa wie bei der verglichenen Art. Stirn mit 



296 v. PUTHZ 

Netzungsspuren, Pronotum und Elytren zwischen den Punkten glatt, Abdomen mit 
kaum erkennbar flacher Netzung. 

Bemerkungen: Diese neue Art unterscheidet sich von ihren Nah verwandten 
durch ihre kurzen, trapezoiden Elytren, ihr vergleichsweise kurzes Pronotum und die 
Sexualcharaktere. Sie ähnelt am meisten den neuguineensischen Spezies S. cyaneoto- 
gatus Puthz und S. magnificus L. Benick. 

Etymologie: Der Namen der neuen Art bezeichnet ihre Herkunft von der 
Molukkeninsel Tanimbar. 

Stenus (Hemistenus) schwendingeri sp. n. 

Material: S -Holotypus und 2 5 5 -Paratypen: THAILAND: Huay Nam Dang, Mae 
Taeng District, 1400 m, 17.XII.1990, P. Schwendinger (MHNG, 1 PT coli. Puthz). 

Beschreibung: Länge: 2,6-3,4 mm (Vorderkörperlänge: 1,6-1,7 mm). Schwarz 
mit leichtem Bronzeschimmer, schwach schimmernd, Vorderkörper grob und sehr 
dicht punktiert, Abdomen vorn ziemlich grob, hinten mäßig fein, überall sehr dicht 
punktiert; Beborstung kurz, wenig auffällig. Fühler schmutziggelb, die Keule gebräunt, 
Kiefertaster rötlichgelb, Beine einfarbig bräunlichgelb. Clypeus schwarz, Oberlippe 
braunschwarz, beide dünn beborstet. Paraglossen oval. 

PM des HT: wH: 28,5; aE: 14; wP: 21; IP: 21,5; wEl: 28,3; LEI: 30; IS: 25. 

Männchen: Beine ohne Auszeichnungen. Metasternum flach gewölbt, grob und 
dicht auf glänzendem Grund punktiert. Vördersternite einfach. 7. Sternit in der hinteren 
Mitte sehr fein und dicht, flach punktiert und beborstet. 8. Sternit mit kleinem 
Apikaiausschnitt etwa im hinteren Achtzehntel. 9. Sternit apikolateral dreizähnig- 
gesägt (Fig. 12). 10. Tergit abgerundet. Aedoeagus (Fig. 13), Medianlobus in 
seiner Vorderhälfte nadeldünn, im Innern mit einem tubigen Innensack und einer 
deutlich skierotisierten Ausstülpspange (Fig. 11). Parameren etwa so lang wie der 
Medianlobus, an ihren Spitzen etwas löffeiförmig erweitert und daselbst mit 3+7 
Borsten, von denen eine besonders lang und kräftig ist. 

Weibchen: 8. Sternit am Hinterrand breit abgerundet. Valvifer apikolateral 
dreizähnig. 10. Tergit abgerundet. Spermatheka aus einem dünnen, vielfach gewunde- 
nen Schlauch ("Schlauchgewirr") mit einem kräftig skierotisierten, ziemlich langen 
Einfüllstutzen bestehend, insgesamt fast so lang wie ein Valvifer. 

Kopf beim Männchen gut so breit wie die Elytren, beim Weibchen etwas 
schmäler, Stirn schmal, eingesenkt und innen ganz flach, Punktierung grob und sehr 
dicht, mittlerer Punktdurchmesser fast so groß wie der apikale Querschnitt des 2. 
Fühlergliedes, Punktzwischenräume viel kleiner als die Punktradien. Fühler schlank, 
zurückgelegt den Pronotumhinterrand überragend, vorletzte Glieder nicht ganz doppelt 
so lang wie breit. Pronotum gut so lang wie breit, in der Vorderhälfte seitlich deutlich 
konvex, hinten deutlich, aber nicht stark, konkav eingeschnürt; keine Eindrücke. 
Punktierung sehr regelmäßig, tief eingestochen, so grob wie auf der Stirn, äußerst 
dicht. Elytren etwa quadratisch, etwas länger als breit, Naht- und Schultereindruck 
flach. Punktierung auffallend regelmäßig, grob und sehr dicht, kaum gröber als am 
Pronotum. Abdomen ellipsoid, Seiten deutlich gerundet, Paratergite des 4. Segments 
nicht ganz so breit wie das 2. Fühlerglied, leicht ventrad geneigt und nur mit einzelnen 



NEUE ORIENTALISCHE STENUS ARTEN 297 

Punkten. Basale Querfurchen der ersten Tergite ziemlich tief, 7. Tergit mit deutlichem 
apikalem Hautsaum (die Art ist voll geflügelt). Punktierung vorn fast so grob wie auf 
der Stirn, hinten erheblich feiner, auf den einzelnen Tergiten jeweils vorn gröber als 
hinten; die mittlere Punktgröße der Punkte des 7. Tergits ist so groß wie eine 
Augeninnenrandfacette; die Punktierung ist überall sehr dicht. 10. Tergit spärlich und 
zerstreut punktiert. Beine schlank, Hintertarsen fast drei Viertel schienenlang, ihr 1. 
Glied gut so lang wie die beiden folgenden zusammengenommen, erheblich länger als 
das Klauenglied; das 4. Glied ist sehr dünn, wenig auffällig gelappt. Stirn und 
Pronotum zeigen nur Netzungsspuren, Elytren deutlich genetzt, Abdomen vorn flach, 
hinten deutlicher genetzt. 

Bemerkungen: Diese neue Art ist vermutlich die Schwesterart des S. pallidipes 
Cameron, sie sieht ihm zum Verwechseln ähnlich, auch der Medianlobus (nicht jedoch 
die Parameren) sind sich ähnlich; sie unterscheidet sich von ihm äußerlich nur schwer 
durch ihre dichtere und etwas weniger grobe Punktierung der Stirnmitte und beim 
Männchen zusätzlich durch viel flacher ausgerandetes 8. Sternit (S. pallidipes zeigt 
hier einen tiefen spitzwinkligen Ausschnitt gut im hinteren Drittel). Unter den 
makropteren, ungemakelten orientalischen (Hemistenus-) Arten mit apikolateral 
gesägtem 9. Sternum und eingesenkter Stirnmitte, die eine ähnliche Größe haben, äh- 
nelt die neue Art noch S. iustus Puthz, dessen Stirnmitte aber gröber punktiert und 
dessen vorderes Abdomen ungenetzt ist, sowie S. humicola Puthz, der jedoch kleinere 
Elytren besitzt. Sie ähnelt auch S. absurdus Puthz und S. peratus Cameron, die aber 
beide eine erhobene Stirnmitte zeigen, ganz abgesehen von den Sexualcharakteren. 

Etymologie: Mit ihrem Namen ehre ich ihren verdienstvollen Sammler, Herrn P. 
Schwendinger (Genf). 

Stenus (Hemistenus) kurbatovi sp. n. 

Material: c?-Holotypus und 4 6 (J-Paratypen: BURMA: E Mandalay, env. Maymyo, 700 
m, 5.m. 1996, Kurbatov (MHNG, 1 PT coli. Puthz). 

Beschreibung: Länge: 4,5-6,0 mm (Vorderkörperlänge: 2,5 mm). Schwarz, 
matt, jede Elytre mit einem ziemlich großen, ovalen, orangeroten Fleck, Stirn grob und 
sehr dicht punktiert, Pronotum sehr grob und sehr dicht, zum Teil zusammenfließend 
skulptiert, Elytren sehr grob, äußerst dicht, rau, kurz-zusammenfließend skulptiert, 
Abdomen vorn grob und dicht, hinten wenig fein und ziemlich dicht punktiert; Be- 
borstung kurz, wenig auffällig. Fühler mittelbraun, die Keule wenig dunkler, 
Kiefertaster rötlichgelb, Beine hellbraun, Schenkel an den Knien wenig dunkler. 
Clypeus schwarz, Oberlippe dunkelbraun, Clypeus ziemlich dicht, Oberlippe dünn und 
spärlich beborstet. Paraglossen oval. 

PM des HT: wH:48; aE: 25; wP: 31; IP: 32,5; wEl: 47; 1E1: 42; IS: 31. 

Männchen: Beine ohne Auszeichnungen. Metasternum gewölbt, wenig fein und 
ziemlich dicht auf flach genetztem Grund punktiert, Hinterhüftumrandung innen glatt. 
Vordersternite einfach. 7. Sternit in der hinteren Mitte feiner und dichter als an den 
Seiten punktiert und beborstet. 8. Sternit mit stumpfwinkligem Ausschnitt etwa im hin- 
teren Fünfzehntel. 9. Sternit apikolateral mit spitzem Zahn. 10. Tergit breit abgerundet. 
Aedoeagus (Fig. 16), Medianlobus vorn in eine breit abgerundete Spitze ver- 
schmälert, im Innern mit einem breittubigen, distal mit schmalem Rohr versehenen 



298 v. PUTHZ 

Innensack und mit einem kleinen, stark skierotisieren distalen Ausstülphaken. 
Parameren wenig länger als der Medianlobus, mit etwa 17+7 langen Borsten. 

Weibchen: unbekannt. 

Bemerkungen: Diese neue Art ist die Schwesterart des S. dentellus L. Benick, 
mit dem sie auch zusammen erbeutet wurde. Sie ähnelt ihm äußerlich zum Ver- 
wechseln, besitzt jedoch einen breiteren Kopf und einen größeren Aedoeagus mit 
anderem Innenbau sowie viel stärker beborstete Parameren. Eine sichere Unter- 
scheidung der beiden Arten dürfte nur mit Hilfe der Genitaluntersuchung möglich sein. 

Etymologie: Ich dediziere diese neue Art ihrem verdienstvollen Sammler, Herrn 
S. A. Kurbatov (Moskau). 

Stenus (Hemistenus) croceipennis sp. n. 

Material: c?-Holotypus und 5 Sa-, 9 9 2-Paratypen: [Indonesien]: Sumatra: Jambi, 
Mt. Kerinci, 1750- 1850 m, IL- 12.XI.1989, Agosti, Lobi, Burckhardt # 11 (MHNG, 5 
Paratypen coll. Puthz). 

Beschreibung: Länge: 4,4-5,6 mm (Vorderkörperlänge: 2,3-2,5 mm). 
Brachypter, schwarz mit bräunlichem Einschlag und schwachem Messingschimmer, 
Elytren mit sehr großem, ovalem, gelblichem Fleck, Stirn grob und sehr dicht punk- 
tiert, Pronotum sehr grob, tief und sehr eng, leicht zusammenfließend punktiert, 
Elytren sehr grob und sehr dicht, etwas längs-zusammenfließend punktiert, Abdomen 
mäßig fein bis sehr fein, fast überall dicht punktiert; Beborstung kurz, kaum auffällig. 
Fühler rötlichgelb, die Keule verdunkelt, Kiefertaster gelb bis rötlichgelb, Beine 
rötlichgelb, die Schenkelspitzen und die Tarsengliedspitzen etwas angedunkelt. 
Clypeus schwarz, Oberlippe schwarzbraun, heller gesäumt, wenig dicht beborstet. 
Paraglossen koniform. 

PM des HT: wH:41; aE: 24; wP: 32; IP: 35,5; wEl: 39; 1E1: 37; IS: 26. 

Männchen: Beine ohne Auszeichnungen. Metasternum dreieckig eingedrückt, 
mäßig grob, dicht auf genetztem Grund punktiert und beborstet, Hinterhüftumrandung 
innen mit wenigen, sehr feinen Punkten. Vordersternite ohne Auszeichnungen. 7. 
Sternit in der hinteren Mitte abgeflacht, sehr fein und dichter als an den Seiten punk- 
tiert und beborstet, Hinterrand kaum erkennbar flach ausgerandet. 8. Sternit mit 
rundlichem Ausschnitt etwa im hinteren Elftel. 9. Sternit apikolateral mit spitzem 
Zahn. 10. Tergit breit abgerundet. Aedoeagus (Fig. 18), Medianlobus vorn 
mäßig breit abgerundet und im Innern außer einem stark skierotisierten, breittubigen 
Innensack ohne weitere, stärker skierotisierte Elemente. Parameren erheblich länger 
als der Medianlobus, apikal lang-löffelförmig und faltig und daselbst mit etwa 19 
mäßig langen Borsten versehen. 

Weibchen: 8. Sternit zur Hinterrandmitte deutlich vorgezogen und daselbst 
abgerundet. Valvifer apikolateral mit spitzem Zahn. 10. Tergit breit abgerundet. Die 
Spermatheka besteht aus einem langen, vierfach gewundenen Schlauch und ist insge- 
samt etwa so lang wie der Valvifer. 

Kopf gut so breit wie die Elytren, Stirn sehr breit mit breiten, flachen 
Längsfurchen, der undeutlich abgesetzte Mittelteil ist so breit wie jedes der 
Seitenstücke, breitrund erhoben und erreicht gut das Augeninnenrandniveau. Die 
Punktierung ist grob und sehr dicht, der mittlere Punktdurchmesser erreicht den 



NEUE ORIENTALISCHE STENUS ARTEN 299 

größten Querschnitt des 3. Fühlergliedes, die scharfen Punktzwischenräume sind viel 
kleiner als die Punktradien, nur auf dem hinteren Mittelteil können sie bis punktgroß 
werden. Fühler schlank, zurückgelegt den Pronotumhinterrand erreichend, vorletzte 
Glieder gut doppelt so lang wie breit. Pronotum wenig länger als breit, in den vorderen 
drei Fünfteln seitlich stark konvex, hinten deutlich konkav eingezogen; die Oberseite 
zeigt mehrere Unebenheiten: eine vorn und hinten abgekürzte mittlere Längsfurche, 
einen kräftigen seitlichen Quereindruck knapp hinter der Mitte und einen flachen 
Quereindruck hinter dem Vorderrand. Die Skulptur ist sehr grob und sehr dicht, unter- 
schiedlich kurz-zusammenfließend, der mittlere Punktdurchmesser liegt über dem 
größten Querschnitt des 3. Fühlergliedes, erreicht aber nicht immer (vor allem direkt 
in der Mitte nicht) den apikalen Querschnitt des 2. Fühlergliedes, die flach genetzten 
Punktzwischenräume sind fast überall kleiner als die Punktradien. Die trapezoiden 
Elytren sind wenig schmäler als der Kopf, deutlich breiter als lang, ihre Seiten hinter 
den wenig ausgeprägten Schultern kräftig erweitert, ihr Hinterrand ist tiefrund aus- 
gerandet. Ihre Oberfläche ist etwas uneben, und zwar erscheinen bei den meisten 
Stücken die großen, gelben Flecken etwas aufgetrieben. Die Flecken können vorn fast 
bis zur Schulter reichen, hinten berühren sie den Hinterrand, seitlich enden sie am 
Deckenabfall und von der Naht sind sie in der Elytrenhinterhälfte etwa um die Länge 
des 2. Hintertarsengliedes getrennt. Die Punktierung ist sehr grob, fließt leicht längs 
zusammen, überall sehr dicht, ähnlich dicht wie am Pronotum. Abdomen breit 
elliptisch, Paratergite breit und leicht aufgebogen, diejenigen des 4. Segments so breit 
wie die Hinterschienen in ihrem basalen Drittel; sie sind mäßig fein, dicht, auch 
nebeneinander punktiert. Die basalen Querfurchen der ersten Tergite sind tief, das 7. 
Tergit trägt einen schmalen apikalen Hautsaum (trotz Brachypterie). Die Tergit- 
punktierung ist vorn mäßig fein und dicht, hinten sehr fein und ziemlich dicht, auf dem 
7. Tergit sind die Punkte kleiner als eine innere Augenfacette, ihre Abstände fast 
doppelt so groß wie die Punkte, das 10. Tergit ist sehr fein und zerstreut punktiert. 
Beine schlank, Hintertarsen etwa zwei Drittel schienenlang, ihr 1. Glied etwas länger 
als die beiden folgenden zusammengenommen, deutlich länger als das Klauenglied; 
das 4. Glied ist lang und schmal gelappt. Die Oberseite des Vorderkörpers ist sehr flach, 
die des Abdomens dicht und deutlich genetzt. 

Bemerkungen: Die neue Art unterscheidet sich von denjenigen (Hemistenus-) 
Arten mit koniformen Paragalossen (vgl. Puthz, 1998: 397 f.), apikolateral spitzem 9. 
Sternum und kurzen, gemakelten Elytren so: von S. raffle si Rougemont durch gröbere 
Punktierung des Vorderkörpers und viel weniger gedrängte Abdominalpunktierung, von 
S. abdominalis maculosus L. Benick, S. cham Puthz (der ein ähnlich punktiertes Ab- 
domen besitzt), kurzflügligen S. coronatus L. Benick, S. leileri Puthz und S. malaba- 
rensis Cameron durch größere Elytrenmakeln, von allen durch die Sexualcharaktere. 

Etymologie: Wegen ihrer großen gelben Elytrenmakeln nenne ich diese Art 
croceipennis. 

Stenus (Hemistenus) agostii sp. n. 

Material: d-Holotypus und 5 S c?-Paratypen: [Indonesien]: Sumatra: Jambi, Mt. 
Kerinci, 3000 m, 12.XI.1989, Agosti, Lobi & Burckhardt # 13a; 3 SS-, 2 9 9-Paratypen: ibi- 
dem, 2500 m, 12.XI.1989, idem # 14 ; 2 SS-, 3 Î-Paratypen: ibidem 1800-2100 m, 6.-7- 
III. 1991, Bocâk & Bocâkova (Naturhistorisches Museum Basel und coli. Puthz). 



300 V. PUTHZ 

Beschreibung: Länge: 4,0-5,4 mm (Vorderkörperlänge: 2,3-2,5 mm). 
Brachypter, schwarz mit schwachem Messingschimmer, Stirn grob und sehr dicht 
punktiert, Pronotum grob, unregelmäßig, zum Teil zusammenfließend punktiert, 
Elytren sehr grob und zusammenfließend skulptiert, Abdomen vorn mäßig grob und 
sehr dicht, hinten fein und dicht punktiert; Beborstung anliegend, wenig auffällig. 
Fühler rötlichgelb, die Keule gebräunt, Kiefertaster gelb, das 3. Glied verdunkelt, 
Beine hell rötlichbraun, die Schenkelspitzen (kaum) und die Tarsen dunkler. Clypeus 
schwarz, Oberlippe braunschwarz, beide wenig dicht beborstet. Paraglossen koniform. 

PM des HT: wH: 40,5; aE: 23; wP: 30,5; IP: 32; wEl: 38,5; 1E1: 39; IS: 31. 

Männchen: Beine ohne Auszeichnungen. Metasternum breit eingedrückt und 
daselbst mäßig grob, dicht auf genetztem Grund punktiert und beborstet, Hinter- 
hüftumrandung innen fast glatt. 3-6. Sternit median unpunktiert, dicht und flach 
genetzt. 7. Sternit median leicht abgeflacht, feiner und viel dichter als an den Seiten 
punktiert und beborstet. 8. Sternit mit schmalem, rundem Ausschnitt etwa im hinteren 
Achtel. 9. Sternit apikal gesägt. 10. Tergit breit abgerundet. Aedoeagus 
(Fig. 19), der schmale Medianlobus zur Spitze breit-dreieckig verengt; neben einem 
stark skierotisierten Innensack wird noch eine wenig stark skierotisierte Aus- 
stülpspange erkennbar. Parameren viel länger als der Medianlobus, an ihren Spitzen 
lang-löffelförmig und daselbst mit zahlreichen kurzen und kräftigen Borsten versehen. 

Weibchen: 8. Sternit am Hinterrand abgerundet, zur Hinterrandmitte kaum 
vorgezogen. Valvifer apikal gesägt. 10. Tergit breit abgerundet. Die Spermatheka 
besteht aus einem doppelt S-förmig gewundenen kräftigen Schlauch mit kleinem 
Blasenanhang und ist etwa zwei Drittel so lang wie der Valvifer. 

Kopf deutlich etwas breiter als die Elytren, Stirn ziemlich breit mit flachen 
Längsfurchen und einem nur schwach erhobenen Mittelteil, der nicht die Höhe der 
Augeninnenränder erreicht. Punktierung grob und sehr dicht, mittlerer Punkt- 
durchmesser so groß wie der mittlere Querschnitt des 3. Fühlergliedes, die scharfen 
Punktzwischenräume deutlich kleiner als die Punktradien, in der hinteren Stirnmitte 
manchmal bis punktgroß. Fühler schlank, zurückgelegt knapp den Pronotumhinterrand 
überragend, vorletzte Glieder etwas länger als breit. Pronotum wenig länger als breit, 
knapp hinter der Mitte am breitesten, von dort nach vorn konvex, nach hinten deutlich 
konkav verengt; außer einer flachen, vorn und hinten abgekürzten (und in der Mitte 
manchmal etwas verbreiterten) mittleren Längsfurche sind noch mäßig flache Schräg- 
eindrücke knapp hinter der Mitte zu sehen. Die Skulptur ist unregelmäßig, oft kurz 
zusammenfließend, die Punkte sind unterschiedlich tief eingestochen, ihre Zwischen- 
räume unterschiedlich stark erhoben, die gröbsten Punkte sind etwas gröber als die 
Stirnpunkte, in der Scheibenmitte stehen meist etwas feinere Punkte. Elytren schmäler 
als der Kopf, etwas länger als breit, Schultern mäßig eckig, Seiten lang erweitert 
(Gesamteindruck: lang-trapezoid), Schultereindruck deutlich, hintere Elytrenmitte 
leicht aufgetrieben. Die Skulptur ist, besonders auf der Hinterhälfte, noch unregel- 
mäßiger und hier auch länger-zusammenfließend als am Pronotum und meist auch 
etwas gröber als dort. Das breit-elliptische Abdomen besitzt breite, dorsad aufgebo- 
gene Paratergite, diejenigen des 4. Segments sind gut so breit die die Hinterschienen 
an ihrer Basis, sie sind dicht, wenig fein, auch nebeneinander punktiert. Die basalen 
Querfurchen der ersten Tergite sind tief, das 7. Tergit trägt einen breiten apikalen 



NEUE ORIENTALISCHE STENUS ARTEN 301 

Hautsaum (obwohl die Art brachypter ist). Die Punktierung ist überall sehr dicht (vorn) 
bis dicht (hinten), die Punkte des 3. Tergits sind so groß wie der basale Querschnitt des 
3. Fühlergliedes, diejenigen des 7. Tergits etwas feiner als eine Augeninnenrandfacette, 
ihre Abstände hier gut punktgroß. Das 10. Tergit ist ziemlich fein und weitläufig punk- 
tiert. An den kräftigen Beinen sind die Hintertarsen nicht ganz zwei Drittel schienen- 
lang, ihr 1. Glied ist deutlich länger als die beiden folgenden zusammengenommen, 
viel länger als das Klauenglied; das 4. Glied ist lang und schmal gelappt. Die Oberseite 
des Vorderkörpers zeigt nur flache Netzungsspuren, das ganze Abdomen ist flach, aber 
deutlich genetzt. 

Bemerkungen: Die neue Art ist die Schwesterart des philippinischen S. 
contrusus L. Benick; von ihm unterscheidet sie sich durch breiteren Kopf, flachere 
Stirnmitte, feiner und dichter punktiertes Abdomen und das gesägte 9. Sternum. Die 
Aedoeagen beider Arten sind sich sehr ähnlich. 

Etymologie: Ich dediziere diese neue Art einem ihrer Sammler, Herrn D. Agosti 
(Zürich). 

DANKSAGUNG 

Folgenden Personen und Instituten sei dafür gedankt, dass sie mir das behan- 
delte Material zur Verfügung gestellt haben: Dr. D. Burckhardt (Naturhistorisches 
Museum, Basel), Dr. G. Cuccodoro und Dr. I. Löbl (Muséum d'histoire naturelle, 
Genf) und Dr. W. Schawaller (Staatliches Museum für Naturkunde, Stuttgart). 

LITERATUR 

Joly, C. 2003. Problématique de l'Entomologie indonésienne, sa complexité. Lambillonea 53: 
155-161. 

Puthz, V. 1971. Revision of the Sfónws-species of New Guinea. Part I. (Coleoptera: 
Staphylinidae). Pacific Insects 13: 447-469. 

Puthz, V. 1980. Die Stenus- Arten {Stenus s.str. + Nestus REY) der Orientalis: Bestimmungs- 
tabelle und Neubeschreibungen (Coleoptera, Staphylinidae). 178. Beitrag zur Kenntnis 
der Steninen. Mit 15 Figuren. Reichenbachia 18: 23-41. 

Puthz, V. 1988. Die Steninen von Borneo II (Coleoptera, Staphylinidae). Neue und alte Arten, 
vorwiegend aus den Ausbeuten des Genfer Naturhistorischen Museums. 218. Beitrag zur 
Kenntnis der Steninen. Mit 1 Tabelle und 45 Textfiguren. Revue suisse de Zoologie. 95: 
631-679. 

Puthz, V. 1991. Über indo-australische Steninen II (Insecta, Coleoptera, Staphylinidae) 222. 
Beitrag zur Kenntnis der Steninen. Mit 48 Figuren. 251. Beitrag zur Kenntnis der 
Steninen. Entomologische Abhandlungen Staatliches Museum für Tierkunde, Dresden 
54: 1-46. 

Puthz, V. 1998. Die Gattung Stenus Latreille in Vietnam (Coleoptera, Staphylinidae). Revue 
suisse de Zoologie 105: 383-394. 



Revision der paläarktischen Arten 
der Gattung Brachygluta Thomson, 1859, 

I.Teil 

(COLEOPTERA StAPHYLINIDAE) 




Giorgio Sabella, Christoph Buckle, Volker Brachat & Claude Besuchet 



Muséum d'histoire naturelle Genève 
2004 



Giorgio SABELLA, Christoph BUCKLE, Volker BRACHAT & Claude BESUCHET 



Revision der paläarktischen Arten der 
Gattung Brachygluta Thomson, 1859, 

1. Teil: Arten mit sekundären männlichen 

Geschlechtsmerkmalen auf den Abdominaltergiten 

(Coleoptera, Staphylinidae, Pselaphinae). 



The systematic position of the genus Brachygluta within the Pselaphinae and 
Brachyglutini is specified. The distribution of the genus covers mainly the 
Holarctig region, with two species known from the Afrotropical region. Detailed 
descriptions and 669 illustrations of the morphology of the external and internal 
skeleton structures as well as of the aedeagus are given for each species. Some 
particular of integument structure and pilosity are reproduced by scanning electron 
microscope photographs. Available information on biology, ecology and onto- 
genesis is summarized. The external morphology and the characteristics of the 
aedeagus are precisely described for the 75 Palaearctic (sub)species exhibiting 
secondary male sexual characters on abdominal tergites. Distribution maps are 
given for all species. 

Twenty-one species are described as new for science, four synonymies are 
established, six subspecies are elevated at species level, and, finally, two new 
combinations are proposed. A list of species and their synonyms is provided. The 
descriptions are in German, the identification keys are in French and English. 

This publication is addressed to systematicians, ecologists, entomologists 
interested in faunistic research as well as to libraries of museums of natural history 
and entomological institutes. 



Instrumenta Biodiversitatis n° 6 

Ed. Muséum d'histoire naturelle, Ville de Genève, March 2004 
283 pages, 669 figures and 15 maps 

ISBN 2-88139-010-2 

Price: 100 Swiss francs + handling and postage 



Orders should be mailed to: Muséum d'histoire naturelle. Publications, 

case postale 6434, CH-1211 Genève 6, Switzerland. 

Fax: +41 22 418 63 01, or to e-mail: publications.mhn@ville-ge.ch 



We are ordering copy(ies) of "Revision der paläarktischen Arten der Gattung 

Brachygluta Thomson, 1859. 1. Teil" 



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Revue suisse de Zoologie 111 (2): 303-307; juin 2004 



Nesticella marapu sp. n., a blind nesticid (Araneae: Nesticidae) 
from Sumba, Indonesia 

Suresh P. BENJAMIN 

Department of Integrative Biology, Section of Conservation Biology (NLU), 

University of Basel, St. Johanns-Vorstadt 10, CH-4056 Basel, Switzerland. 

Present address: University of California, Berkeley, Insect Biology Division - ESPM, 
201 Wellman Hall #3112 Berkeley, CA 94720-3112, USA. 
E-mail: Sureshb@nature.berkeley.edu 



Nesticella marapu sp. n., a blind nesticid (Araneae: Nesticidae) from 
Sumba, Indonesia. - A new species of the genus Nesticella is described 
from both sexes. It is characterised by the presence of a beak-shaped para- 
cymbium. It has long legs and weak pigmentation but lacks eyes. 

Key-words: Nesticella - Nesticidae - new species - troglobit - caves. 

INTRODUCTION 

The spider family Nesticidae has been revised by Lehtinen & Saaristo (1980). 
It is the sister taxon of Theridiidae (Coddington, 1986; Coddington, 1989; Coddington, 
1990; Griswold et al., 1998). Spiders of both families construct webs with a supporting 
structure and with lines under tension termed gumfooted lines. The bases of these gum- 
footed lines are the only parts of the web that contain viscid silk (Benjamin & 
Zschokke, 2002; Benjamin & Zschokke, 2003). Nesticids are mostly found in caves. 
Among nesticid spiders collected during an expedition to the island of Sumba by the 
Naturhistorisches Museum, Basel, two adult specimens of an undescribed species were 
discovered. This new species has long legs, is pale and lack eyes, typical morpho- 
logical adaptations for life in caves. Details of the expedition and a description of the 
locality are given in Bühler & Sutter (1951). 

MATERIAL AND METHODS 

Morphological structures were examined on temporary mounts embedded in 
glycerin. All drawings were made with a Nikon Labophot-2 and a Nikon SMZ-U 
microscope with drawing tube. The methods are described in detail in Benjamin (2000) 
and in Benjamin & Jocqué (2000). Measurements are in mm. All specimens examined 
are deposited in the Naturhistorisches Museum, Basel, NMB. Terminology of male 
palpal sclerite follows Lehtinen & Saaristo (1980) and Coddington (1990). 



Manuscript accepted 13.01.2004 



304 



S. P. BENJAMIN 



TAXONOMY 

Nesticeila marapu sp. n. Figs 1-5 

Types: S holotype (NMB 2751a) and ? paratype (NMB 2751b), Indonesia, Sumba, 
Weimangura, Marro Cave, 440m, 25.08.1949, leg. Alfred Bühler and Ernst Sutter. 

Etymology: The specific name is derived from Marapu, meaning spirits of ancestors. The 
main aim of the traditional Sumbanese religion is to maintain a peaceful and fruitful relationship 
with Marapu. Used as a noun in apposition. 

Diagnosis: Nesticella marapu sp. n. can be distinguished from all Nesticella 
species by the beak-shaped paracymbium and by the shape of the conductor. 

Description: Male (holotype). Colouration pale yellow. Carapace flattened, 
covered with some erect setae, mostly confined to the cephalic area. Eyes absent, no 
traces of eye sockets visible. Sternum covered with erect setae. Chelicerae without 
teeth, fangs reddish. Legs densely covered with thin long setae. Abdomen with some 
faint grey markings, covered with erect setae. Colulus very small, much smaller than 
that of the female, two erect setae present. Palpal paracymbium (PC) bifurcate, beak- 
shaped, both branches with distal modifications (Figs 1-2). Median apophysis (MA) 
tapering to a point. Tegular apophysis (TA) present in the form of a blunt elevation 
(Fig. 1; corresponding to "tegular apophysis 1" in Lehtinen & Saaristo (1980) and 
"theridiid tegular apophysis" of Coddington (1990). 





1 



Figs 1-2 
Nesticella marapu sp. n. 1. Left male palp, ventral view. 2. Ditto, retrolateral view. C conductor; 
E embolus; MA median apophysis; PC paracymbium; TA tegular apophysis. Scale line: 0.2 mm. 



NESTICELLA MARAPU SP. N. 



305 




Figs 3-5 
Nesticella marapu sp. n. 3. Epigynum, ventral view. 4. Vulva, ventral view. 5. Ditto, dorsal view. 
CD copulatory duct; CO copulatory opening; FD fertilization duct; S spermatheca; SC scape; SP 
sclerotised plate. Scale line: 0.2 mm. 



306 S. P. BENJAMIN 



Measurements: total length 2.3; carapace length 1.2; carapace width 1.1. Legs 



II III IV 



femur 


2.8 


1.7 


1.3 


patella 


0.4 


0.4 


0.4 


tibia 


2.5 


2.2 


1.8 


metatarsus 


2.4 


1.0 


0.6 


tarsus 


1.0 


0.9 


0.5 



2.0 

0.4 
2.3 
1.7 
0.8 



total 9.1 6.2 4.6 7.2 

Female. Carapace bright yellow, flattened, covered with erect setae. Eyes 
absent, no traces of eye sockets visible. Sternum bright yellow, with many erect setae. 
Chelicerae yellow with three reddish teeth on promargin of the groove, fangs reddish, 
sickle-shaped. Legs pale yellow, slender, densely covered with long setae. Palp with a 
long, sickle-shaped claw. Abdomen pale yellow with diffuse grey markings, covered 
with erect red-brown setae. Colulus large, with two setae on ventral surface. 

Epigynum with a square scape projecting outwards (SC in Fig. 3) and extending 
into a thin sclerotized plate beyond the scape (SP in Fig. 4). Copulatory openings (CO) 
originating at lateral sides of the scape. Copulatory ducts (CD) leading inwards to the 
spermathecae (S). Fertilization ducts (FD) as in Fig. 5. 

Measurements: total length 2.6; carapace length 1.2; carapace width 1.0. Legs: 

I II III IV 

1.3 2.0 

0.3 0.3 

1.0 1.8 

1.0 1.8 

0.7 0.7 

total 7.0 5.2 4.3 6.6 

Distribution: Known only from the type locality. The Marro cave is about 3 km 
long and situated close to Kabalidana, west of Waimangura (Bühler & Sutter, 1951). 

ACKNOWLEDGEMENTS 

I am indebted to Ambros Hänggi (NMB) for providing the interesting speci- 
mens. Brigitte Braschler (Basel) and Peter Schwendinger (Geneva) are thanked for 
critical review of the manuscript. This study was supported by the Swiss National 
Science Foundation (Grant no. 31-55617.98 to Samuel Zschokke) and the by Uni- 
versity of Basel. 

REFERENCES 

Benjamin, S. P. 2000. Epidius pan'ati sp. n., a new species of the genus Epidius from Sri Lanka 
(Araneae: Thomisidae). Bulletin of the British Arachnological Society 11: 284-288. 



femur 


2.0 


1.7 


patella 


0.3 


0.3 


tibia 


1.7 


1.0 


metatarsus 


2.2 


1.5 


tarsus 


0.8 


0.7 



NESTICELLA MARAPU SP. N. 307 



Benjamin, S. P. & Jocqué, R. 2000. Two new species of the genus Suffasia from Sri Lanka 
(Araneae: Zodariidae). Revue suisse de Zoologie 107: 97-106. 

Benjamin, S. P. & Zschokke, S. 2002. Untangling the tangle-web: web building behaviour of 
the comb-footed spider Steatoda triangulosa and comments on phylogenetic implica- 
tions (Araneae: Theridiidae). Journal of Insect Behavior 15: 791-808. 

Benjamin, S. P. & Zschokke, S. 2003. Webs of theridiid spiders: construction, structure and evo- 
lution. Biological Journal of the Linnean Society 78: 293-305. 

Bühler, A. & Sutter, E. 1951. Sumba-Expedition des Museums für Völkerkunde und des 
Naturhistorischen Museums in Basel, 1949. Verhandlungen der Naturforschenden 
Gesellschaft in Basel 65: 181-217. 

Coddington, J. A. 1986. The monophyletic origin of the orb web (pp. 319-363). In: Shear, W. 
A. (ed.). Spiders - Webs, Behavior, and Evolution. Stanford University Press, Stanford. 

Coddington, J. A. 1989. Spinneret silk morphology: evidence for the monophyly of orb- 
weaving spiders, Cyrtophorinae (Araneidae), and the group Theridiidae plus Nesticidae. 
Journal of Arachnology 17: 71-95. 

Coddington, J. A. 1990. Ontogeny and homology in the male palpus of orb- weaving spiders and 
their relatives, with comments on phylogeny (Araneoclada: Aranoidea, Deinopoidea). 
Smithsonian Contributions to Zoology 496: 1-52. 

Griswold, C. E., Coddington, J. A., Hormiga, G. & Scharef, N. 1998. Phylogeny of the orb- 
web building spiders (Araneae, Orbiculariae: Deinopoidea, Araneoidea). Zoological 
Journal of the Linnean Society 123: 1-99. 

Lehtinen, P. T. & Saaristo, M. I. 1980. Spiders of the Oriental-Australian region, II. Nesticidae. 
Annales Zoologici Fennici 17: 47-66. 



Revue suisse de Zoologie 111 (2): 309-313; juin 2004 



Dikerogammarus villosus (Crustacea: Amphipoda): 
another invasive species in Lake Geneva 

Loïc BOLLACHE 

Equipe Ecologie Evolutive, UMR CNRS 5561 Biogéosciences, Université de 

Bourgogne, 6 Blvd Gabriel, F-21000 Dijon, France. 

E-mail: bollache@u-bourgogne.fr 



Dikerogammarus villosus (Crustacea: Amphipoda): another invasive 
species in Lake Geneva. - According to field observations performed in 
2002 and 2003, the amphipod species Dikerogammarus villosus (Sowinsky, 
1 894) is now well established in Lake Geneva. The author predicts that this 
and future invasions by D. villosus will have serious effects on freshwater 
ecosystems. 

Key-words: Dikerogammarus villosus - amphipod - biological invasion - 
ecological impact. 

INTRODUCTION 

The Ponto-Caspian amphipod (crustacean) Dikerogammarus villosus (Sowin- 
sky, 1894) is currently among the most successful invaders of aquatic ecosystems in 
Europe (Dick & Platvoet, 2000; Bij de Vaate et al, 2002). Being a native of the Ponto- 
Caspian Region, this species is widely distributed in the Black Sea, Azov Sea and 
Caspian Sea basins, from the lower reaches and delta of the Danube River to the Volga 
River (Carausu et al., 1955). After invading the Danube in Austria in 1989, D. villosus 
was collected for the first time in the upper reaches of the river Danube in 1992 
(Neseman et al., 1995). In 1994 the species was recorded in the lower Rhine River (Bij 
de Vaate & Klink, 1995). Two years later D. villosus was discovered in the Moselle 
River and in 1997 for the first time in the French hydrosystem in the Saone River 
(Devin et al, 2001; Bollache, 2003). In 1998 it was recorded in the Rhone River and 
in the Meuse River. Two years later D. villosus was detected in the Seine River (in 
2000) and three years later in the Loire River (in 2001). Field surveys and laboratory 
experiments indicate that the ecological impact of a D. villosus invasion may be severe 
(Van der Velde et ai, 2000). For example, Dick and Platvoet (2000) have shown in 
Netherland fresh waters that D. villosus was rapidly eliminating Gammarus duebeni 
(Lilljeborg, 1851) and Gammarus tigrinus (Sexton, 1939), respectively, a native and an 
invader species from North America. In addition, D. villosus, being larger than the 
other freshwater amphipod species in western Europe, is known to prey on numerous 
macroinvertebrates (Dick et al., 2002), and an isotope analysis (ô 15 N) indicated that it 



Manuscript accepted 30.10.2003 



310 



L. BOLLACHE 



is active at the same trophic level as fish species (Marguillier, 1998). My study, based 
on field observations in 2002 and 2003, demonstrates that D. villosus is now 
established in Lake Geneva. 

MATERIAL AND METHODS 

Four sites in France were sampled (Fig. 1). Yvoire, Corzent, Thonon-les-Bains 
and Lugrin. Gammarids were collected by kick-sampling using a hand net. D. villosus 
was identified by using the key of Carausu et al. (1955). Only adult individuals were 



Lausanne 



Morges 



Vevey 

Montreux 




Versoi 



Genève 



Fig. 1 
Map of Lake Geneva with sampling sites (full triangles), and number of D. villosus specimens 
collected at each site on certain dates. 



examined, as most of the species-specific diagnostic characters are fully developed and 
more obvious in these specimens. Dikerogammarus can easily be separated from 
Gammarus by the striking difference in the morphology of their third uropods. In the 
genus Gammarus the inner and outer rami are well developed, with the inner ramus 
varying from 1/3 to 3/5 of the length of the outer ramus (Fig. 2a), while in the genus 
Dikerogammarus the inner rami are vestigial (Fig. 2b). D. villosus males and females 
bear two protuberances with spines on the back of the first and second urosome (see 
fig. 3), and short setae on the peduncle of the second antennae. Moreover there is 
strong sexual dimorphism in the flagellum of the second antennae, which is distally 
much more densely setose in males than in females (Fig. 2c). Long and dense setae are 
also present on the first and second gnathopods of males (Fig. 2d). 



DIKEROGAMMARUS VILLOSUS 



311 




outer ramus 



"'"H 

Fig. 2 
(a) Gammarus pulex, third uropod after Roux, 1967; (b) Dikerogammarus villosus, third uropod 
after Carausu et al, 1955; (c) D. villosus male and female, second antenna after Carausu et al, 
1955; (d) D. villosus male, first and second gnathopods after Carausu et ah, 1955. 



RESULTS 

D. villosus was observed for the first time in Lake Geneva during the summer 
of 2002 at Lugrin (France); one specimen was found among specimens of the native 
gammarid, Gammarus fossarum. In 2003 D. villosus was recorded at the other three 
French sites prospected, Yvoire harbour, Thonon-les-Bains (INRA station) and Conzert 
harbour (Fig. 3), without any other gammarid species present and was found to be very 
abundant in rockpools and under porous stones, hanging on tightly to the bumps on the 
stones. 



DISCUSSION 

The present and previous data allow us to attribute the arrival of D. villosus to 
Lake Geneva between the years 2001 and 2002. The reason for this recent massive 
invasion of a Ponto-Caspian species in central and western Europe is the increasing 
ionic content of large European rivers in the last decades, caused by industrial and agri- 
cultural pollution (Jazdzewski & Konopacka, 2002). Another cause is explained in the 
"invasional meltdown hypothesis" (Ricciardi & Rasmussen, 1998; Simberloff & Von 
Holle, 1999; Ricciardi, 2001) according to which a previously invasion of the zebra 
mussel Dreissena polymorpha (Pallas, 1771) in the central and western European 
freshwater ecosystem may have facilitated the invasion of D. villosus by providing 
substrate, food and shelters. The same scenario is invoked to explain the establishment 
and dispersal of Chaeiogammaris ischnus (Stebbing, 1906) another Ponto-Caspian 
amphipod, in the Great Lakes of North America (Witt et al, 1997; Dermott et al., 1998; 



312 L. BOLLACHE 




Fig. 3 
Dikerogammarus villosus adult male after Carausu et al., 1955, modified after Bollache, 2003. 



Nalepa et al., 2001). Dick et al. (2002) have recently shown in the Netherlands that 
prédation by the invader D. villosus was significantly greater than prédation by the 
native amphipod species G. duebeni. The predatory capacity of D. villosus, together 
with its high ability to disperse by anthropogenic means, lead I to predict that D. 
villosus will soon be present in the others lakes of Switzerland, and that this poses a 
serious threat for the biodiversity of freshwater ecosystems in Europe. 

ACKNOWLEDGMENTS 

The author thanks Dr B. Lods-Crozet and Dr C. Gillet for their help in obtaining 
documents on Lake Geneva and in collecting some individuals of D. villosus. 

REFERENCES 

Bu De Vaate, A., Jazdzewski, K., Ketelaars, H. A. M., Gollash, S. & Van der Velde, G. 
2002. Geographical patterns in range extension of Ponto-Caspian macroinvertebrate 
species in Europe. Canadian Journal of Fisheries and Aquatic Sciences 59: 1159-1174. 

Bu De Vaate, A. & Klink, A. G. 1995. Dikerogammarus villosus Sowinsky (Crustacea: 
Gammaridae) a new immigrant in the Dutch part of the Lower Rhine. Lauterbornia 20: 
51-54. 

Bollache, L. 2003. Trois espèces d'Amphipodes épigés nouvelles pour la Saône bour- 
guignonne. Bulletin Scientifique de Bourgogne 51: 1-9. 

Carausu, S., Dobreanu, E. & Manolache. C. 1955. Crustacea Amphipoda, forme salmastre si 
de apa dulce. In: Bodnariuc, N. et al. (eds). Fauna repubicci populäre romîne 4, 
Crustacea 4. Bucaresti, 409 pp. 

Dermott, R., Witt, J., Um, Y. M. & Gonzalez, M. 1998. Distribution of the Ponto-Caspian 
amphipod Echino gammar us ischnus in the Great Lakes and replacement of native 
Gammarus fasciatus. Journal of Great Lakes Research 24: 442-452. 

Devtn, S., Beisel, J. N., Bachmann, V & Moreteau, J. C. 2001. Dikerogammarus villosus 
(Amphipoda: Gammaridae): another invasive species newly established in the Moselle 
river and French hydrosystems. Annales de Limnologie 37: 21-27. 



DIKEROGAMMARUS VILLOSUS 3 1 3 



Dick, J. T. A. & Platvoet, D. 2000. Invading predatory crustacean Dikerogammarus villotWm- 

inates both native and exotic species. Proceedings of the Royal Society of London B 267: 

977-983. 
Dick, J. T. A., Platvoet, D. & Kelly, D. W. 2002. Predatory impact of the freshwater invader 

Dikerogammarus villosus (Crustacean: Amphipoda). Canadian Journal of Fisheries and 

Aquatic Sciences 59: 1078-1084. 
Jazdzewski, K. & Konopacka, A. 2002. Invasive Ponto-Caspian species in waters of the Vistula 

and Oder basins and the southern Baltic sea (pp 384-398). In: Leppäkoski, E., 

Gollasch, S. & Olenin, S (eds). Invasive aquatic species of Europe. Distribution, 

impacts and management. Kluwer Academic Publishers, Dordrecht, Boston, London. 
Marguillier, S. 1998. Stable isotope ratios and food web structure of aquatic ecosystems. PhD 

thesis, University of Brussels. 
Nalepa, T. F., Schloesser, D. W., Pothoven, S. A., Hondorp, D. W., Fanslow, D. L., 

Tuchman, M. L. & Fleischer, G. W. 2001. First finding of the amphipod Echino- 

gammarus ischnus and the mussel Dreissena bugensis in Lake Michingan. Journal of 

Great Lakes Research 27: 384-391. 
Nesemann, H., Pockl, M. & Wittmann, K. J. 1995. Distribution of epigean Malacostraca in the 

middle and upper Danube (Hungary, Austria, German). Miscellanea Zoologica 

Hungarica: 10 49-68. 
Ricciardi, A. 2001. Facilitate interactions among aquatic invaders: is an 'invasional meltdown' 

occuring in the Great Lakes? Canadian Journal of Fisheries and Aquatic Sciences 59: 

1144-1158. 
Ricciardi, A. & Rasmussen, J. B. 1998. Predicting the identity and impact of future biological 

invaders: a priority for aquatic ressource management. Canadian Journal of Fisheries 

and Aquatic Sciences 55: 1759-1765. 
Roux, A. L. 1967. Les gammares du groupe pulex. Essai de systématique biologique. Thèse, 

Faculté des Sciences de l'Université de Lyon, 172 pp. 
Simberloff, D. & Von Holle, B. 1999. Positive interactions of nonindigenous species: invasion 

meltdown? Biological Invasions 1: 21-32. 
Van der Velde, G., Rajagopal, S., Kelleher, B., Musko, I. & Bu de Vaate, A. 2000. 

Ecological impact of crustacean invaders: general considerations and examples from the 

Rhine river (pp. 3-34). In: von Vaupel Klein, J. C. & Schräm, F. R. (eds). Proceedings 

of the fourth International Crustacean Congress, Amsterdam, The Nederlands, July 20- 

24, 1998, vol 2. Rotterdam, The Nederlands: Balkema. 
Witt, J. D. S., Hebert, P. D. N. & Morton, W. B. 1997. Echino gommar us ischnus: another 

crustacean invader in the Laurentian Great Lakes basin. Canadian Journal of Fisheries 

and Aquatic Sciences 54: 264-268. 



Revue suisse de Zoologie 111 (2): 315-369; juin 2004 



Revision of the European athecate hydroids and their medusae 

(Hydrozoa, Cnidaria): 

Families Oceanidae and Pachycordylidae 

Peter SCHUCHERT 

Muséum d'histoire naturelle, CP 6434, CH-1211 Genève 6, Switzerland. 

E-mail: Peter.Schuchert@mhn.ville-ge.ch 



Revision of the European athecate hydroids and their medusae 
(Hydrozoa, Cnidaria): Families Oceanidae and Pachycordylidae. - This 
paper reviews the European Hydrozoa species belonging to the families 
Oceanidae and Pachycordylidae. Emended diagnoses for the Oceanidae, the 
Pachycordylidae, and for some of their genera are provided. Due to the 
paucity of suitable characters, the macrotaxonomy of the two families is un- 
settled and must be regarded as provisional. The family Oceanidae includes 
the genera Cordylophora, Corydendrium, Merona, Oceania, Rhizogeton, 
Turritopsis, Turritopsoides, and the new genus Similomerona gen. n. The 
family Pachycordylidae includes the genera Pachycordyle and Silhouetta. 
Turritopsis polycirrha and T dohrnii are both regarded as valid species and 
distinct from T. nutricula of the western Atlantic. Histological sections 
confirmed that Turritopsis polycirrha is hermaphroditic and larviparous, 
while T. nutricula is dioecious and oviparous. Turritopsis rubra (Farquhar, 
1895) comb. n. from New Zealand is also regarded as a valid species and 
removed from synonymy with T nutricula. The known distribution of 
Corydendrium dispar is extended to include the Faroe Islands. A former 
record of C. dispar from Denmark is referred to Pachycordyle navis 
(Millard, 1959) comb. n. Pachycordyle fusca Müller, 1913 is regarded as a 
likely synonym of Pachycordyle pusilla (Motz-Kossowska, 1905) comb. n. 

Key-words: Marine invertebrates - Cnidaria - Hydrozoa - Antoathecata - 
Oceanidae - Pachycordylidae - Clavidae - descriptions - revision - taxo- 
nomy. 

INTRODUCTION 

Although the athecate hydroids and anthomedusae of the European coasts have 
a long taxonomic history, with some species going back to Linnaeus, we lack a 
comprehensive, up-to-date treatise like the one by Cornelius (1995) for the thecate 
hydroids of North- West Europe. Some groups are covered by good monographs. The 
Stylasteridae have been monographed by Zibrowius & Cairns (1992). The European 
Hydra species are treated by Holstein (1995). The monograph of Brinckmann-Voss 



Manuscript accepted 04.09.2003 



316 P. SCHUCHERT 

(1970) covers the Capitata, but is restricted to the Mediterranean fauna only. Naumov's 
(1969) book on the Hydrozoa of the USSR includes also the Baltic Sea, but its scope 
is not European for the most part. Russell's (1953) work on the British medusae is 
certainly still very valuable, but treats only species with a medusa phase. There are also 
some recent worldwide reviews of single families which also include European 
species, e. g. by Segonzac & Vervoort (1995) for the genus Candelabrum, Boero et al. 
(2000) for Zanclea, or Schuchert (2001b) for the Corynidae. Some similar, older 
publications, e. g. Vannucci & Rees (1961), Rees (1962), also offer much valuable 
information. However, the scattered information makes it desirable to review all 
species. The author therefore aims at bringing together in a series of publications all 
important taxonomic information on the European Anthoathecata. 

All European species belonging to the order Anthoathecata (Cornelius, 1995) 
will be revised, as will the supraspecific taxa where necessary. Taxa for which there 
exist recent revisions will, however, not be treated in detail. Wherever possible, taxa 
will be redefined so that most of them are plausibly monophyletic. However, these 
animals offer only few morphological characters and they tend to be insufficiently 
complex. Many shared characters are thus very prone to be homoplasies. A compre- 
hensive phylogenetic analysis based exclusively on morphology was therefore 
considered unreliable and only a few suitable characters could be used to delimit some 
genera and families. 

The geographic region covered in this revision reaches from the North Cape to 
the Canary Islands and the European parts of the Mediterranean. It excludes the Azores 
and Iceland. The hydroid species of the Azores are listed in Rees & White (1966) and 
Cornelius (1992). The hydroid fauna of Iceland has recently been reviewed by 
Schuchert (2001a). 

In the present paper, the families Oceanidae and Pachycordylidae have been 
revised. The macrotaxonomy of both families, as well as their genera, is particularly 
unsettled and disputed. It can therefore only be regarded as provisional. 

MATERIAL AND METHODS 

Standard histological techniques were used to obtain serial thin sections. 
Animals fixed in formaldehyde were embedded in paraffin and serial sections of 8-10 
pirn thickness were made. Staining was with Hematoxilin and Eosin. All slides have 
been deposited in the MHNG. 

Abbreviations 

BMNH The Natural History Museum, London, England 

ICZN International Code of Zoological Nomenclature, 4 th Edition 

IRSNB Institut Royal des Sciences Naturelles de Belgique, Bruxelles, Belgium 

MHNG Muséum d'histoire naturelle de Genève, Switzerland 

RMNH Naturalis, Leiden, The Netherlands 

ZMUC Zoological Museum Copenhagen, Denmark 

ZSM Zoologische Staats Sammlung, Munich, Germany 



OCEANIDAE AND PACHYCORDYLIDAE 3 1 7 

TAXONOMIC PART 

Oceanidae Eschscholtz, 1829 

Synonyms - Cordylophoridae von Lendenfeld, 1885. 

Diagnosis - Anthoathecata Filifera with monomorphic or polymorphic polyps. 
Gastrozooids bearing filiform tentacles scattered over at least distal third of hydranth 
body, not concentrated in a narrow band. Gonophores developing either below the 
hydranth body, or directly on stolons, or from stolonal blastostyles with perisarc- 
covered pedicels. Gonophores either free medusae or fixed sporosacs. Free medusa, 
where present, with four mouth lips, mouth margin fringed with numerous spherical 
nematocyst clusters; four simple radial canals, without centripetal canals, juveniles 
with four or more tentacles, adults with numerous tentacles, these not grouped, their 
nematocysts evenly distributed. Ocelli present on adaxial bases of tentacles. Gonads on 
interradial walls of manubrium. 

Remarks - The name Oceanidae Eschscholtz, 1829 is here revived and it 
replaces the junior synonym Cordylophoridae von Lendenfeld, 1885 as well as the 
name traditionally used for the present group: Clavidae McCrady, 1859. 

If the genus Oceania Péron & Lesueur, 1810 is regarded as valid, then auto- 
matically also the family name Oceanidae Eschscholtz, 1829 becomes available. If the 
genera Oceania and Cordylophora are placed within the same family, then Oceania is 
a senior synonym of Cordylophoridae von Lendenfeld, 1885. The latter name was 
recently re-introduced by myself (Schuchert 2001a, 2003a, 2003b) without being 
aware of the name Oceanidae Eschscholtz, 1829. 

The scope of the family Oceanidae as used here corresponds largely to what was 
until recently known as Clavidae (comp. Millard, 1975; Bouillon, 1985; Calder, 1988a; 
Schuchert, 1996; Bouillon & Boero, 2000). The possession of scattered filiform 
tentacles was usually considered a diagnostic trait of the family Clavidae. However, 
also other filiferan families, like the Pandeidae or Hydractiniidae, have polyps with 
scattered tentacles. The hydroid of Stomotoca atra L. Agassiz, 1862, a member of the 
family Pandeidae, has scattered filiform tentacles (Boero & Bouillon, 1989) and in the 
absence of information on the medusa phase would certainly have been classified 
among the former Clavidae. Scattered tentacles are also prevalent in the sister group of 
the Filifera, the Capitata (see Petersen, 1990 for diagnoses for both taxa). The character 
"scattered filiform tentacles" must thus be used with necessary caution, although it is 
potentially a synapomorphy for the family Oceanidae. The isolated occurrences of 
scattered filiform tentacles in the Pandeidae and Hydractiniidae must then be explained 
as homoplasies. 

The genus Clava does not belong to the Oceanidae. Schuchert (2001a) showed 
that the genus Clava Gmelin, 1791 must be transferred to the family Hydractiniidae. 
Although Clava multicornis (Forsskâl, 1775), the type species of Clava, has scattered 
tentacles similar to many Oceanidae, its gonophores develop on the hydranth body 
directly below the tentacles. This gonophore position is the plesiomorphic condition 
within the Anthoathecata and it is found in filiferan families such as the Hydractiniidae 
and Eudendriidae, as well as in many other hydroids, e. g. Corynidae, most Zancleidae, 
some Limnomedusae, and some Leptothecata lacking a gonotheca. In some Filifera, 



318 P. SCHUCHERT 

however, the site of gonophore development has been shifted towards proximal, either 
to the hydranth pedicels, branches, or even stolons. This downward shift is an apo- 
morphy found in the Oceanidae, Bougainvilliidae, Pandeidae, Cytaeididae and some 
other minor groups. In the Calycopsidae, the site of the gonophores is somewhat 
unclear because they are produced at the insertion of the naked pedicel with the 
hydranth body (comp. Brickmann-Voss, 1979; Schuchert, 1996). 

Because the genus Clava shows the plesiomorphic condition of gonophore 
position, and because there are no evident synapomorphies, it cannot be included in the 
Oceanidae as defined here. Clava is better transferred to the Hydractiniidae. Some 
Hydractiniidae have similarly scattered filiform tentacles (comp. Millard, 1975; 
Schuchert, 1996; Bouillon et al. 1997) and there are actually no significant morpho- 
logical differences that would distinguish Clava multicornis from the Hydractiniidae, 
particularly when compared to Clavactinia. The similarity of Clava with the 
Hydractiniidae has also been noted by Boero et al. (1998), but they regarded it as 
distinct on account of its monomorphic polyps. This is, however, not convincing as 
also several hydractiniids lack a polymorphism and gonophores develop on gastro- 
zooids that remain otherwise unchanged (e. g. Hydractinia sarsii; see Schuchert, 
2001a). Gonophore producing hydranths generally reduce their size and tentacle num- 
bers, an effect named reproductive exhaustion. This reproductive exhaustion should 
not be confounded with a true, pre-determined polymorphism, in which gonozooids 
differ from gastrozooids even before they develop gonophores. 

Because Clava, the genus to which Clavidae is anchored, must be transferred to 
the Hydractiniidae, the taxon name Clavidae is not available anymore for the 
remaining genera of this family (Clavidae actually becomes a senior synonym of 
Hydractiniidae, see Schuchert 2001a). The name Oceanidae is therefore proposed for 
these remaining genera. 

The original scope of Oceanidae Eschscholtz, 1829 was very broad and encom- 
passed genera now allocated to several families. This is certainly due to the original 
broad meaning of the genus Oceania, but this should not be seen as an argument 
against its usage. Many other old hydrozoan names that are still in use have a similar 
history (e. g. the Linnean genera Tubularia and Sertularia). The family Oceanidae is 
tied to the genus Oceania and applies to the family that contains this genus, irres- 
pective of the original scope of the family. The historical usage of Oceania is 
summarized under remarks for the genus Oceania. 

The family Oceanidae thus comprises the following previously known genera: 
Cordylophora, Corydendrium, Merona, Oceania, Rhizogeton, Turritopsis, and Turrit- 
opsoides. Additionally, the new genus Similomerona is proposed. 

The family Oceanidae as defined here lacks a clear synapomophy, hence its 
delimitation from other families like the Bougainvilliidae and Calycopsidae is not strict 
(Millard, 1975; Calder, 1988a expressed similar thoughts). It is possible that the family 
Oceanidae is not a monophyletic group and future results of sequence phylogenies 
might require a regrouping. The taxonomy of the Oceanidae must thus be regarded as 
provisional only. 

It was necessary to re-define some of the genera included in this family, both to 
accommodate some recently described species which do not entirely fit formerly 



OCEANIDAE AND PACHYCORDYLIDAE 319 

provided diagnoses and to use as much phylogenetic information as possible to delimit 
monophyletic groups. Nearly all species described as belonging to these genera were 
considered for the diagnoses, although not all are discussed here. Unfortunately, the 
available synapomorphies are very limited and I failed to produce a convincing phylo- 
genetic analysis. The genera were therefore delimited in a way that minimizes name 
changes and maximizes to some degree the possible monophyly. The synapomorphies 
and some important problems of the genera are discussed in the following sections. 

Considering only the life-cycle, species with a medusa phase like Turritopsis 
nutricula or Oceania armata must represent the least derived condition because a 
medusa was present in the ground plan of the Anthoathecata (see Petersen, 1990; 
Schuchert, 1993; Boero et al, 1998). 

The current usage of Turritopsis is unmistakable and the genus is well deli- 
mited. The vacuolated gastrodermal cells forming a peduncle on top of the manubrium 
represents a clear apomorphy. 

The genus Oceania, although in current usage quite well defined, poses taxo- 
nomic problems. They are outlined under remarks for the genus. Oceania auct. is only 
gradually different form Turritopsis and both genera should perhaps be merged, as it 
has been done by Haeckel (1879). Also McCrady (1857) thought that Turritopsis is 
very closely related to Oceania auct., and he initially considered Turritopsis to be only 
a subgenus of Oceania. To maintain nomeclatural stability, both genera are here 
employed. Traditionally, Turritopsis was kept separate from Oceania auct. on account 
of its manubrial peduncle formed by vacuolated cells (Mayer, 1910; Kramp, 1961). 
Oceania has a gelatinous peduncle, but the most proximal parts of the radial canals are 
also dilated and composed of large cells, exactly as in Turritopsis. In Oceania and in 
large Turritopsis medusae, these vacuolated cells are continued along the manubrium 
as four perradial ribs, resembling claws that clasp the manubrium (Figs 2C, 3B, 5). The 
vacuolated cells and the radial ribs could be a synapomorphy of the two genera. 
Oceania differs from Turritopsis in having stalked nematocyst clusters along it mouth, 
while they are without a distinct stalk in Turritopsis. Adult Turritopsis and Oceania 
medusae also resemble medusae of the family Pandeidae (comp. Kramp, 1959), but 
the lining of the mouth margin with nematocysts clusters separates them from the latter 
family (see Fig. 5C). These nematocyst clusters are, however, not a unique synapo- 
morphy, as also at least some medusae of the family Calycopsidae have them (see e. g. 
Bythotiara spec, in Schuchert, 1996: fig. 9e). Medusae of the family Calycopsidae are, 
however, easily separable on account of their tentacle morphology. 

Corydendrium parasiticum, type species of its genus, has gastrodermal gonads 
developing within the perisarc tubes of the hydranth pedicels and they are only 
extruded when mature. This is a clear apomorphy. The same internal gonads are also 
found in Corydendrium brevicaulis Hirohito, 1988, although this species forms 
variably either branched or stolonal colonies. Corydendrium parasiticum possesses 
another apomorphy: the side-branches remain within the perisarc tube of the original 
branch and they emerge only after a certain distance (Weismann, 1883). The perisarc 
of the side-branch is already formed within the tube of the original branch, leading to 
a typical nested structure of the perisarc tubes, best seen in cross-sections (well visible 
even in hand-made sections). This clear apomorphy is also found in Corydendrium 



320 P. SCHUCHERT 

dispar Kramp, 1935 (see Kramp, 1935: fig. 2). However, C. dispar has external 
gonophores, although the structure of the male ones resemble that of C. parasiticum. 
Turritopsis nutricula hydroids also form erect, branching colonies with adnate side- 
branches. The perisarc tubes of Turritopsis, however, are not nested. This was verified 
in hand-made cross-sections of colonies from the Mediterranean and the Andaman Sea. 
The adnate side branches of T. nutricula represent perhaps only the first steps of a 
development that lead to the nested tubes seen in Corydendrium parasiticum. Because 
Turritopsis does not possess any of the two apomorphies found in Corydendrium, 
Kramp 's (1935) proposal of merging Turritopsis and Corydendrium is here not 
adopted. This would also cause very troublesome name changes. 

Merona has very obvious apomorphies: the nematophores in perisarc tubes and 
the sporosacs on blastostyles. The genus thus corresponds very likely to a clade. 
Antsulevich & Polteva (1986) described a new Merona species from the Kurile Islands 
which they named Merona nematophorum. It has naked nematophores resembling 
modified hydranths and it produces sporosacs issuing directly from the stolons. These 
combined differences suggest that M. nematophorum has perhaps attained its super- 
ficial similarity to other Merona species by convergence. Unfortunately, all these 
animals offer too few morphological characters to substantiate this with the aid of a 
reliable phylogenetic analysis and we have to wait for a molecular phylogenetic inves- 
tigation. Meanwhile, it seems appropriate to follow Calder's (1988a) opinion that M. 
nematophorum should be transferred to a new genus. Therefore, the new genus Similo- 
merona proposed. 

The diagnosis for Similomerona gen. n. is as follows: "Anthoathecata Filifera 
with polymorphic polyps; feeding zooids stolonal, with scattered filiform tentacles; 
defensive zooids with rudimentary tentacles, not in perisarc tubes; gonophores 
sporosac developing on stolons." Type species is Merona nematophorum Antsulevich 
& Polteva, 1986. The species name will thus become Similomerona nematophorum 
(Antsulevich & Polteva, 1986) comb. n. The genus is currently monotypic. It is not 
represented in the European fauna. 

Rhizogeton, with its sporosacs on the stolons and the stolonal or sessile 
hydranths, is easily distinguished from other members of the family. Both characters 
are apomorphies. but they are not independent and the Rhizogeton-like morphology 
can be imagined to have easily originated from various other genera. It is thus very 
prone to be polyphyletic. 

Turritopsoides brehmeri Calder, 1988b, the type species of the genus, normally 
forms purely stolonal colonies, but infrequently some shoots bear a side-branch. These 
side-branches are adnate to the branch of origin, an apomorphy also found in Turri- 
topsis nutricula. This, and the branched spadix of the female sporosacs, justify keeping 
the genus separate from Rhizogeton. The genus Turritopsoides is not represented in the 
European fauna. 

The genus Cordylophora is difficult to define in terms of evident apomorphies, 
and the delimitation to Pachycordyle, here placed in a separate family, is based prima- 
rily on the tentacle arrangement of the hydroids (for an opposing view see Moni, 1980; 
or Stepanjants et al., 2000). In the Pachycordylidae, the tentacles are confined to a 
small band below the hypostome, while in the Oceanidae they are more scattered. 



OCEANIDAE AND PACHYCORDYLIDAE 321 

Some genera that have at times been assigned to the Clavidae sensu Bouillon 
(1985) are either invalid or do not belong to the Oceanidae as defined here: 

Campaniclava Allman, 1864: referred to the Pandeidae (Picard, 1956). 

Clava Gmelin, 1791: see above and Schuchert (2001a), referred to Hydractiniidae. 

Cnidostoma Vanhòffen, 1911: this genus was referred to the Clavidae sensu Bouillon, 
1985 by Picard (Picard in Kramp, 1961: 444), a proposal accepted by Bouillon 
(1985). Kramp (1959, 1961) allocated it to the Cytaeididae. Examination of 
Cnidostoma fallax from the Ivory Coast (MHNG INVE 27133) confirmed the 
accurate description of Vanhòffen (1911). The overall habit, but especially the 
manubrium ending in four clusters of stalked nematocytes, bring this species 
very close to medusae of the Hydractiniidae and Cnidostoma is here provi- 
sionally allocated to that family. 

Hataia Hirai & Yamada, 1965 is excluded from the Oceanidae because its sole species 
{Hataia parva) possesses stenoteles and must thus be transferred to the Capitata 
(see Yamada & Kubota, 1991). 

Tubiclava Allman, 1863: Allman (1872: 256) wrote: "...since I do not at this distance 
of time feel absolutely certain as to the identity of the Dublin Bay and Torquay 
hydroids, the gonosome of Tubiclava must still be regarded as in some respects 
hypothetical, and the genus itself as defined above must be accepted with just 
so much of a provisional element." The type species of the genus Tubiclava is 
T. lucerna (by monotypy). Allman (1863, 1872) described Tubiclava lucerna 
based on infertile material. He thought that he had observed similar colonies a 
few years earlier that had gonophores, but which he previously thought to be an 
aberrant Clava multicornis. Rees (1956: 505) maintained that Tubiclava lucerna 
is a problematic species and he doubted whether Allman had actually observed 
sporosacs. Tubiclava is a doubtful genus and should therefore not be used 
anymore. Although occasionally other authors mention findings of Tubiclava 
lucerna, this species remains doubtful and no material could be located in any 
museum. All other species allocated to Tubiclava are either not recognizable or 
must be referred to other genera. Tubiclava fruticosa Allman, 1 872 was based 
on infertile material and is not recognizable. Picard (1958: 189), referred all 
Mediterranean records of T. fruticosa to Pachycordyle napolitana, but some in 
Riedl (1959) are evidently Turritopsis dohrnii. Also Tubiclava rubra Farquhar, 
1895 belongs to the genus Turritopsis (see Schuchert, 1996). Tubiclava laxa 
Fraser, 1938 must be referred to the genus Merona, while Tubiclava triserialis 
Fraser, 1938 is not recognizable. In his diagnosis of Tubiclava Allman (1872) 
stated that the sporosacs develop on the hydranth body immediately below the 
tentacles, just as in Clava multicornis. This renders the genus incompatible with 
the concept of the Oceanidae as used here. The theoretical genus Tubiclava 
sensu Allman has some resemblance to Millardiana Wedler & Larson, 1986. 
Millardiana longitentaculata Wedler & Larson, 1986, presently the only species 
of its genus, is not known from the region under investigation, but Cornelius 
(1992) reported it from the Azores and it is conceivable that this species could 
also be found along the coasts of southern Europe. 



322 P- SCHUCHERT 

Mìllardìana Wedler & Larson, 1986 has sporosacs on the hydranth body. Therefore, 
using the same arguments as given above for Clava, it can neither be included 
in the Oceanidae nor the Bougainvilliidae as proposed by Calder (1988a). At 
present, the affinities of Millardiana are unclear. It should preferably be placed 
in a new family. 

Pachycordyle Weismann, 1883: this genus is here placed in a separate family, the 
Pachycordylidae. For more details see under remarks for this family (includes 
also Thielia Stepanjants et al., 2000). 

Rhìzodendrium Calder, 1988a: it might be necessary to use this genus for some stolonal 
Oceanidae to separate them from Rhizogeton fiisiformis L. Agassiz, 1862 (type 
species of the genus). It is not used here because we do not yet know the 
gonophores of the type species of Rhìzodendrium (see under genus Rhizogeton). 

The order of the genera given below has no phylogenetic meaning. It merely 
attempts to group resembling genera to facilitate comparisons. 

Turritopsis McCrady, 1857 

Synonyms - Clavula Wright, 1859; Dendroclava Weismann, 1883. 
Type species - Turritopsis nutricula McCrady, 1857. 

Diagnosis - Erect, branching hydroid colonies, side-branches and hydranth 
pedicels adnate to for some distance, pensare tubes not nested. Hydranths spindle- to 
club-shaped; filiform tentacles scattered over much of hydranth body. Gonophores 
develop on the hydrocauli in perisarc covered region and are liberated as medusae. 
Medusae with proximal portion of radial canals swollen through vacuolated gastro- 
dermals cells, forming a peduncle-like mass on top of manubrium, in larger animals 
continued on manubrium as perradial ribs. Tentacles numerous and evenly distributed. 
Mouth four-lipped, fringed with numerous spherical nematocyst clusters, these without 
distinct stalk. Ocelli adaxial, at tentacle base. 

Remarks - For the taxonomic history of this genus see Calder (1988a). For the 
publication date of McCrady 's works see Calder et al. (1992). 

Large Turritopsis and Oceania medusae resemble each other quite closely. Both 
have large cells at the proximal end of their radial canals which are continued along the 
manubrium as four perradial ribs, resembling claws that clasp the manubrium (Figs 2C, 
3B, 5). Oceania differs from Turritopsis in having stalked nematocyst clusters along it 
mouth, while they are without a distinct stalk in Turritopsis. In addition, the manu- 
brium base of Oceania armata is somewhat constricted. This allows to distinguish 
even badly preserved material (Kramp, 1965). 

Turritopsis polycirrha (Keferstein, 1862) Figs 1 & 2C-D 

? Medusa sanguinolenta Modeer, 1791: 26. 

? Cyanea coccinea Davis, 1841: 236, pi. 2 figs 1-3. 

? Oceania globulosa Forbes, 1848: 29, pi. 3 fig. 3. 

Turris neglecta - Wright, 1859: 106; Hincks, 1868: 13, pi. 3 fig. 1. 

[not Turris neglecta Lesson, 1843]. 

Clavula Gossii Wright, 1859: 106. pi. 8 fig. 1. 

Oceania polycirrha Keferstein, 1862: 26, pi. 2 figs 11-13. 



OCEANIDAE AND PACHYCORDYLIDAE 



323 




'*MAAMAA* aA 



Fig. 1 
Turritopsis polycirrha (Keferstein, 1862), sketch after preserved material from The Netherlands, 
note that the radial canals overtop the mass of vacuolated cells, a few eggs are visible on the 
lower part of the manubrium, tentacles are lost, scale equals 1 mm. 



? Tubiclava fruticosa Allman, 1872: 257, pi. 2 figs 5-6. 

Turritopsis polynema Haeckel, 1879: 66, new name. 

Turritopsis polycirrha - Hartlaub, 1897: 480, pi. 16c fig 2; Hartlaub, 1911: 202, figs 179-180. 

Turritopsis nutricula - Kramp, 1930: 13, map 5; Russell, 1953: 115, figs 54A-C, 55, 56, pi. 5 

figs 1-5, pi. 29. 
[not Turritopsis nutricula McCrady, 1857]. 

Material of T polycirrha - ZMUC, as T nutricula, North Sea, 50° 59' N 1° 37'5 E, collected 
23 August 1904, about 5 medusae, some with planulae, bells not so well preserved, one 
used for serial sectioning, deposited as MHNG INVE 33967 - RMNH Coel 10345, as T. 
nutricula, Texel, The Netherlands, North Sea, coll. 1973; 19 medusae, 18 clearly with 
oocytes or embryos, two specimens with visible oocytes were used to make serial 
sections, deposited as MHNG INVE 34417 and 34418 - IRSNB, hydroids as T. nu- 
tricula, coll. J. Bouillon June 1964 Roscoff, 2 lots, one small colony on microscopic 
slide, overgrowth of Fosliella farinosa indicates that it has been cultivated for quite some 
time; the second lot is a small polyp with gonophores growing on a piece of an uniden- 
tified gelatinous animal. 

Material of T. nutricula for comparisons - ZMUC, Beaufort Inlet, North Carolina, USA, > 
100 medusae, collected August to September 1962 and 1964 by J. Allwein, many mature 
animals, sexes clearly distinguishable, no brooding, 2 females and 2 males used for serial 
sections, deposited as MHNG INVE 33963 to 33966. 

Material of T. rubra for comparisons - see also Schuchert (1996), as T. nutricula - About 50 
living medusae collected around Rangitoto Island (Hauraki Gulf, New Zealand), July 
2002, some deposited as MHNG INVE 33469, 2 males and 2 females used for serial 
sections (MHNG INVE 33968 to 33971). 

Type material - There presumably exists no type material of T. polycirrha. 

Diagnosis - Typical Turritopsis medusae, adults 4-5 mm in height and dia- 
meter, 80-90 tentacles, tentacle tips not inflated, umbrella top rounded, manubrium 
without gelatinous peduncle, on top of manubrium four blocks of vacuolated cells 
fused into a single compact mass, radial canals overtop this mass. Vacuolated cells 
continued downward on manubrium as bulging, perradial rib. Radial canals broad. 
Simultaneous hermaphrodites and larviparous. Colour: stomach and gonads brilliant 
red to dark crimson. Nematocysts of medusa (Russell, 1940): microbasic euryteles 



324 P. SCHUCHERT 

(8-10) x (4) ^m; desmonemes (6-7) x (4-4.5) pirn. Polyp phase inadequately known 
from nature, likely a stolonal or only sparingly branched colony, hydranths with 
scattered filiform tentacles, hydranth colour: red. 

Description - See Russell (1953), who also provides good colour drawings. 

Biology - Neritic species. Medusa present in autumn and winter months, 
occasionally until April; most abundant in September and October (Russell, 1953). 
Along the coast of The Netherlands, Van der Baan (1980) found it in December and 
January only. According to Kramp (1930), the medusae are presumably liberated from 
the hydroid during June or July. Teissier (1965) indicates August as the month of 
medusa liberation. Medusae present in the North Sea are considered to originate from 
the English Channel (Russell, 1953; Edwards, 1968). 

Distribution - English Channel; Great Britain, up to Firth of Forth in the east, 
up to Bristol Channel in the west; southern parts of the North Sea, sometimes as far 
east as Heligoland. Type locality: St. Vaast, Normandy, France. 

Comparison of reproductive strategies - The European Turritopsis poly- 
cirrha and T dohrnii, as well as other nominal Turritopsis species from the Pacific 
have been synonymized with the American T. nutricula (although McCrady (1857) did 
not mention where he had collected T. nutricula, it can be inferred from McCrady 
( 1 859) that Charleston Harbor must be the type locality). Particular attention was there- 
fore paid to Turritopsis medusae from the east coast of the USA and they were 
compared to Turritopsis medusae from other regions. The morphology of more than 
100 medusae originating from the USA were examined. They all matched the 
description of T nutricula given by Mayer (1910) and Brooks (1886). Together with 
the proximity of the type locality, there is no doubt about their specific identification. 
Even cursory examination with a stereomicroscope reveals that the sexes must be 
separate as eggs are easily discernible in mature females. None of the females had 
larvae or developing embryos attached to the manubrium. Histological examination of 
two males and two females confirmed that individuals produced either sperms or eggs, 
but never both together (Figs. 2A-B). The spermatogenic cells are small cells with little 
cytoplasm and a strongly staining nucleus. They differentiate distally into spermato- 
gonia and finally into spermatozoa (comp. Tardent, 1978). In clusters of mature or 
nearly mature sperms, their aligned tails become visible even under light microscopy 
by producing a filamentous structure. The American T nutricula is thus clearly 
dioecious and oviparous. This has been noted by other authors as well (Brooks, 1886; 
Brooks & Rittenhouse, 1907). 

About 24 Turritopsis polycirrha medusae from the eastern Atlantic (North Sea) 
were obtained. No unambiguously identifiable males were found among them, but 
almost all specimens had eggs and some even had attached planulae. Due to the limited 
number of animals, only three medusae could be used for serial sectioning. The histo- 
logical organization of the three proved to be clearly different from that observed in the 
American T. nutricula. Besides oocytes of all developmental stages, there often are 
pockets containing small cells with little cytoplasm that show differentiation into 
typical spermatozoa (Figs. 2C-D). Comparison with male T. nutricula from the USA 



OCEANIDAE AND PACHYCORDYLIDAE 



325 



w 




y:& 



g 




su 




9 i ..• % 



Bfc 






rc 



B 



-. 






^'■i^- 



m 



». 



g 



g. 




/" ^0 



m 






"-^ 



OC 



-s. 

■ 






, . su 

C 



/ 



«._ 



» 



Fig. 2. Horizontal thin sections in region of gonads. A) Male Turritopsis nutricula medusa from 
the USA, arrow points to spermatogenic cells, scale equals 50 firn. B) Female Turritopsis nutri- 
cula medusa from the USA, arrow points to a developing oocyte, scale equals 50 /im. C) 
Turritopsis polycirrha, mature hermaphroditic medusa from the North Sea, overview, arrow 
points to a developing embryo, the gonad contains oocytes as well as pockets of spermatogenic 
cells. Note the perradial vacuolated cells of the gastrodermis. Scale equals 20 //m. D) Higher 
magnification of animal depicted in C, arrowhead points towards spermatogenic cells, arrow 
points to an oocyte, scale equals 50 pirn. Abbreviations: g = gastrodermis, oc = oocyte, rc = radial 
canal, su = subumbrellar musculature. 



326 P. SCHUCHERT 

leaves no doubt that these must be spermatogenic cells as well as mature spermatozoa. 
The presence of eggs, developing embryos, and sperms in various stages of differen- 
tiation makes the European T. polycirrha a simultaneous hermaphrodite. This confirms 
similar observations by Hartlaub (1911), which, however, have largely been ignored. 
The fertilized eggs are brooded and released as fully formed planula larvae, a fact that 
has already been observed by several authors (Mayer, 1910; Russell, 1953). 

Turritopsis medusae from New Zealand, until recently attributed to T. nutricula 
(see Schuchert, 1996), appear to be almost indistinguishable from T. polycirrha (Fig. 
3A-B). Both have the same morphology and colour, and the same morphological 
arguments can be used to distinguish them from T. nutricula (table 1). However, in 
contrast to T. polycirrha, the New Zealandic Turritopsis is gonochoristic. Examination 
of more than 100 living, mature Turritopsis medusae from New Zealand (see also 
Schuchert, 1996) clearly showed that males and females can be distinguished using a 
stereomicroscope (10 x magnification). The females brood their larvae, sometimes 
even to the primary polyp stage (Fig. 3A-B; see also Schuchert, 1996). The exami- 
nation of histological serial sections (2 females and 2 males) confirmed that the two 
sexes are clearly separate and that there are no differentiating sperm cells in the female 
medusae (results not shown). Because Turritopsis from New Zealand is thus distinct 
from both T. nutricula and T polycirrha, it is preferable to regard it as a distinct 
species, namely Turritopsis rubra (Farquhar, 1895) new. comb. 

Taxonomic History - Because the medusa of T polycirrha is quite conspi- 
cuous due to its red colour, it was possibly mentioned by several early naturalists (see 
synonymy above and Russell, 1953) but it was Keferstein (1862) who gave the first 
reliable description. Due to the great similarity to the American Turritopsis nutricula, 
Mayer (1910) considered T. polycirrha to be only a synonym of the former. Kramp 
(1930) and Russell (1953) followed the arguments of Mayer, although Russell 
acknowledged the differences between the European and the American forms. Hartlaub 
(1911, 1917) and Browne (1916), both of whom presumably had seen living European 
material, did not agree and kept T polycirrha separate from T nutricula. Browne 
(1916: 179) kept both species distinct based on the following arguments: 

T nutricula: "... the radial canals, by a considerable thickening of their walls 
and coalescence, form a kind of endodermal peduncle upon which the stom- 
ach hangs." 

T. polycirrha: "... a further development of the endoderm takes place, it is no 
longer confined to the walls of the radial canals, but grows across the top of 
the umbrella cavity and forms a homogenous mass of cells. This extension 
also grows downward and takes in the radial canals in the uppermost part of 
the subumbrella. The extension of the endoderm is recorded by the leaving 
of the outer half of the radial canal wall in situ alongside the umbrella. In 
adult specimens, the radial canals run alongside this mass of endoderm 
nearly to the top of the subumbrella and there they curve sharply inside to 
the stomach." 

Browne's arguments translate into the observation that in T. polycirrha the four 
masses of vacuolated cells are completely fused and the outer portions of the radial 



OCEANIDAE AND PACHYCORDYLIDAE 



327 



canals reach higher up and overtop the mass of vacuolated cells. The difference in the 
radial canals was also observed in the examined preserved material (comp. Figs 1 and 
3D-E). In addition, the American Turritopsis nutricula has its mass of vacuolated cells 
more clearly split into four blocks, while in the European T. polycirrha, the four blocks 
are tightly fused into one mass, although the fusion lines remain visible. Hartlaub 
(1917: 407) also noted this difference. As shown in Fig. 3, in living medusae these 
differences are quite obvious (although Figs 3A-C and 3D-E compare T. rubra and T. 
nutricula, but the former species from New Zealand closely resembles T. polycirrha). 
Fixation and contraction, though, may largely obliterate the morphological differences. 
The observed differences of the three compared species are summarized in table 1 . 

Hartlaub (1911) and Russell (1953) regarded Turris neglecta Lesson, 1843 as a 
questionable synonym of T. polycirrha or T. nutricula. According to Lesson (1843), 
Turris neglecta has a size of about 12-15 mm and only 16 tentacles. This is incom- 
patible with Turritopsis polycirrha and the identity of Turris neglecta remains unre- 
solved. The Turris neglecta described by Gosse (1853), Wright (1959), and Hincks 
(1868), however, are without doubt referable to Turritopsis polycirrha. 



Table 1 : 


: Differences of the 


Turritopsis species 


discussed in this publication. 


character 


T. nutricula 
(eastern Atlantic) 


T. polycirrha 
(North Sea) 


T. rubra 
(New Zealand) 


T. dohrnii 
(Mediterranean) 


embryo brooded 


no 


yes 


yes 


no 


reproduction 


gonochoristic 


hermaphroditic 


gonochoristic 


gonochoristic 


outer portion of 
radial canals 


does not appear to 
overtop mass of 
vacuolated cells 


appears to 
overtop mass of 
vacuolated cells 


appears to 
overtop mass of 
vacuolated cells 


appears to 
overtop mass of 
vacuolated cells 


mass of 
vacuolated cells 


split into four 
blocks 


four blocks fused 
into one mass 


four blocks fused 
into one mass 


split into four 
blocks 


colour of 
stomach 


yellow 
to orange 


red-orange 
to red 


red-orange 
to red 


?brownish 


bell size 


3-4 mm, 
max. 6 mm 


4-5 mm 


3-7 mm 


1.8-2.7 mm 


umbrella top 


rather fiat 


rounded 


rounded to 
conical 


conical 


swelling of 
tentacle tips 


distinctly 
swollen 


not swollen 


sometimes 
slightly swollen 


swollen 


tentacle number 


40-100 


up to 90 


up to 120 


14-32 


manubrial pads 
in young medusae 


not reported 


unknown 


present 


present 


hydroid colony 


branched, mono- 
or polysiphonic 


? stolonal or 

sparingly 

branched 


branched, mono- 
or polysiphonic 


branched, mono- 
or polysiphonic 



328 P. SCHUCHERT 

Remarks on Polyp stage - Although Turritopsis polycirrha medusae can be 
quite common in the English Channel (Russell, 1953, as T. nutricula), the hydroid 
stage has only rarely been reported. Actually, the only records appear to be by Billard 
(1912) and Teissier (1965), who found it at Roscoff. Teissier recorded it on pebbles in 
the intertidal zone and on old shells in subtidal regions. Billard (1912, as T. neglecta) 
found the hydroid in aquaria. He did not mention gonophores. In addition, it is very 
likely that Tubiclava fruticosa Allman, 1872 is the polyp phase of T. polycirrha. 
Especially worth mentioning is its reddish colour, a characteristic trait of several 
Turritopsis hydroids. However, Airman's hydroid from Wales did not bear medusae 
buds and remains of uncertain identity. 

The hydroid has been reared several times from planulae released by the 
medusa (e. g. Gosse, 1853 as Turris neglecta; Wright, 1859 as Clavula Gossi; Russell, 
1953 as T. nutricula). Cultivated hydroids and colonies from nature collected by Prof. 
J. Bouillon near Roscoff (Brittany) were deposited in the IRSNB and were examined 
for this study. However, the state of conservation did not permit extraction of infor- 
mation beyond that indicated above under "characteristics". Nevertheless, Turritopsis 
polycirrha appears to form only small, sparingly branched colonies. It seems quite 
unlikely that T. polycirrha forms large colonies as seen e. g. in T dohrnii or T. rubra. 
Such conspicuous colonies would certainly have been discovered along the coasts of 
Europe, one of the regions of the world that has been searched for hydroids very 
intensively. 

Conclusions - Although quite similar in morphology, Turritopsis medusae 
from the western and north-eastern Atlantic differ in reproductive strategy. While the 
American medusae spawn their eggs into the free water (oviparity), their European 
counterparts brood fertilized eggs and release fully formed planula larvae (larviparity). 
There is also a significant difference in the gonad structure: the European medusae are 
simultaneous hermaphrodites, while American medusae are dioecious. Whether there 
exist also purely male medusae in the European Atlantic cannot be said at present, 
because not enough specimens could be examined. None of the prominent authors (e. 
g. Hartlaub, 1911; Russell, 1953) mentions male medusae, but Keferstein (1862) noted 
that he had observed male and female medusae. It is also not known whether auto- 
fertilization is possible. Hermaphroditic hydromedusae are not common, but the 
phenomenon is known to occur e. g. in Eleutheria dichotoma (see Müller, 1908). 

The brooding behaviour and the hermaphroditism make the Turritopsis 
medusae of the north-eastern Atlantic quite distinct from T. nutricula found in the 
western Atlantic. Although Russell (1953) attributed only varietal character to this 
difference, it seems unlikely that animals with such different reproductive strategies 
form a single biological species. Because there are also other, unrelated differences 
(table 1), it is justified to regard T. polycirrha as a valid species and distinct from T. nu- 
tricula. 

The medusa of Turritopsis polycirrha is very similar to T. rubra from New 
Zealand inasmuch both species brood their embryos, share the same intense red colour, 
the tightly fused blocks of vacuolated cells, and the high insertion point of the radial 
canals (Fig. 3A-B). Turritopsis rubra differs, however, in being gonochoristic and pre- 
sumably also in forming large, polysiphonic hydroid colonies (see also table 1). 



OCEANIDAE AND PACHYCORDYLIDAE 



329 




Fig. 3 
A-C, Turritopsis rubra from New Zealand; this species has almost the same morphology and 
colour as the European T. polycirrha. D-E, Turritopsis nutricala from Brazil (photographs D-E 
kindly provided by Dr A. Migotto, Sâo Paulo). A) Brooding female medusa, bell height 4 mm. 
B) Higher magnification of manubrium with planula larvae. C) Hydroid stage, tentacles some- 
what contracted, height of hydranth body about 1 mm. D-E) Medusa, bell size approx. 3 mm, 
note the difference to T. rubra in bell shape, manubrium colour, shape of the vacuolated masses, 
the swollen tentacle tips, and the entry point of the radial canals. 



Another European Turritopsis species is the Mediterranean T. dohrnii, the status 
of which is discussed below. 

Further potentially valid nominal species within the genus Turritopsis are: 
Turritopsis fascicularis Fraser, 1943; Turritopsis pleurostoma (Péron & Lesueur. 



330 P. SCHUCHERT 

1810); Turritopsis chevalense (Thomely, 1904); Turritopsis lata von Lendenfeld, 
1885; Turritopsis pacifica Maas, 1909; and Turritopsis minor (Nutting, 1905). Their 
reproductive strategies and life-cycles must all be re-evaluated. 

Turritopsis dohrnii (Weismann, 1883) Fig. 4 

Dendroclava dohrnii Weismann, 1883: 26; Zoja, 1892: 409; Neppi, 1917: 42, pi. 4 fig. 14. 

Tubiclava fruticosa - Riedl, 1959: 611. 

[not Tubiclava fruticosa Allman, 1872 = ? Turritopsis polycirrha]. 

Turritopsis nutricula - Neppi & Stiasny, 1913: 25; Ramil & Vervoort, 1992: 17; Piraino et ah, 

1996: 304, figs 4-5; Carla et al. 2003: 213, figs 1-3. 
[not Turritopsis nutricula McCrady, 1857]. 

Material examined - Several living polyp colonies collected at Mallorca, depth 2-4 m, July- 
August 1999 and 2000, initial stages of medusa phase cultivated; some material 
deposited as MHNG INVE 27123 - Living material from Ischia, Italy, collected July 
1993, medusae cultivated for 7 days - Natural History Museum of Vienna, as Tubiclava 
fruticosa, coll. Riedl, Tyrrhenia Expedition, 31.7.1956, on sponge, medusae buds present 
although tissue somewhat deteriorated (material mentioned in Riedl, 1959). 

Type material - There presumably exists no type material of T dohrnii. 

Diagnosis - Hydroid colony of variable height, either sparingly branched with 
shoots only a few mm high to much branched and polysiphonic colonies up to 35 mm 
high. Hydranths with 12-20 tentacles. Periderm two-layered (see Schuchert, 1996: fig. 
5d). Hydranths in life colourless or pinkish. Newly released medusa with 8-12 
tentacles, manubrium with yellow-fluorescent interradial pads (paler to colourless in 
cultivated animals), vacuolated cells visible. Adult medusa (after Neppi & Stiasny, 
1913; Piraino et al., 1996) up to 2.7 mm in height, diameter 3.2 mm, mature specimens 
with 14-32 tentacles, manubrium reaching to bell margin, tentacles sometimes with 
terminal swellings, ocelli rust-coloured, gonads brownish, with 4 interradial rust- 
coloured dots, proximal ends of radial canals swollen through vacuolated gastrodermal 
cells, the four swellings not fused into a single mass. Sexes separate and females 
oviparous (Dr. S. Pirano, pers. comm.). Medusa can metamorphose back into polyp 
stage (Piraino et al, 1996; Carla et al, 2003). 

Distribution - Western Mediterranean, Adriatic Sea. Type locality: Naples, 
Mediterranean. 

Remarks - Weismann (1883) used the name Dendroclava dohrnii for this 
species. The spelling of the specific epithet ending with -ii is correct and must be main- 
tainted (ICZN, 1999: § 31.1.1 and §31.1.3. The name is meant as a dedication to Anton 
Dohrn, latinized as Dohrnius, which gives in genitive case dohrnii). 

The initial description of Turritopsis dohrnii was based on the hydroid phase 
only. It was synonymized with T. nutricula by Neppi (1917) and her opinion was 
adopted by nearly all subsequent authors. The only life-cycle studies resulting in 
mature medusae are described by Piraino et al (1996, as T. nutricula) and Carla et al. 
(2003, as T. nutricula). 

I consider T. dohrnii to be a distinct species differing from the European T. poly- 
cirrha and the American T. nutricula. Turritopsis dohrnii is oviparous and dioecious 
and thus clearly distinct from the hermaphroditic and larviparous T. polycirrha. Other 
differences from the latter species are more gradual: colony size (large polysiphonic 



OCEANIDAE AND PACHYCORDYLIDAE 



331 



SN \W/ 'Z 




Fig. 4 
Turritopsis dohrnii (Weismann, 1883). A) Silhouette of colony, preserved material from 
Mallorca, scale equals 5 mm. B) Hydranth and medusa bud, tentacles somewhat contracted, after 
living material from Ischia, scale bar 0.4 mm. C) Young medusae, drawn after photograph in 
Piraino et al. (1996), scale bar 1 mm. 

versus small monosiphonic), number of tentacles of the medusa (16-32 versus up to 
90), the colour of the gonads (brown versus red), and possibly also the brown spots on 
the gonads in T. dohrnii (the latter detail needs a re-evaluation). 

The differences of T. dohrnii and the American T. nutricula are significant, 
although they are mostly only gradual. The medusae differ in tentacle numbers, bell 
size, and the size of the vacuolated masses. The newly released medusae have conspi- 
cuous manubrial pads, which have never been reported for T. nutricula (comp. Brooks 
1886; Brooks & Rittenhouse, 1907; Migotto, 1996). The polyp stage seems to be 
indistinguishable, although T. dohrnii may form larger colonies. According to Migotto 
(1996), T. nutricula forms either monosiphonic or polysiphonic colonies, as in T. 
dohrnii. It seems prudent to regard T. dohrnii as separate from T. nutricula until more 
detailed life-cycle studies prove otherwise. Preliminary molecular investigations 
confirmed that the two belong to two separate lineages (M. P. Miglietta, Duke 
University, pers. comm.). 



332 P- SCHUCHERT 

While the hydroid phase is relatively common on rocky substrates of the 
western Mediterranean, Turritopsis medusae are much less known. Turritopsis 
medusae have has been reported from the Mediterranean by: Hartlaub (1911, Séte, as 
T. polycirrha), Neppi & Stiasny (1913, Trieste, as T. nutricula), Schmidt (1973, as 
T. nutricula), Dowidar (1984, Egyp, as T. nutricula), Goy et al. (1991, Lebanon, as 
T. nutricula). 

Another record of a Mediterranean Turritopsis medusa is T. pacifica by Browne 
(1927). Browne's medusa from Port Said at the Suez Canal was 3.5 mm in size and had 
120 tentacles, a number which is significantly higher than observed by Neppi & 
Stiasny (1913) or Piraino et al. (1996). I believe that this medusa was probably not T. 
dohrnii and that it was perhaps an invader from the Red Sea or Indian Ocean. Kramp 
(1959: 246) likewise stated that T nutricula might have invaded the eastern Medi- 
terranean via the Suez Canal, but is unclear to what morphotype he was referring to or 
on which evidence he based his assumption. 

The records of T. nutricula by Schmidt (1973), Dowidar (1984), and Goy et al. 
(1991) from the eastern Mediterranean do not provide sufficient data to evaluate the 
taxonomic status of their animals. 

Oceania Péron & Lesueur. 1810 

Type species - Mayer (1910) designated Oceania armata Kölliker, 1853 as type species, a 
species not originally included in the genus Oceania by Péron & Lesueur (1810). 
Mayer's designation is thus currently not valid. 

Diagnosis - Anthoathecate medusae with eight or more marginal tentacles not 
grouped in clusters. Manubrium upon a simple, solid, gelatinous peduncle not formed 
by vacuolated cells. Manubrium ovoid, diameter at base smaller than in middle, mouth 
rim studded with a row of spherical nematocyst clusters, each cluster on a short stalk. 
Ocelli adaxial, on tentacle base. Polyp phase forming stolonal colonies, hydranths with 
scattered filiform tentacles. 

Remarks - Oceania Péron & Lesueur, 1810 originally included species of 
medusae now referred to several different families and orders (Goy, 1995). First 
attempts to restrict the scope of the genus Oceania were made by Lesson (1843), who 
used it more in the sense of Clytia of present-day authors, thus falling within the order 
Leptothecata. Forbes (1848). on the contrary, used Oceania so that its species would 
today either fall within the Pandeidae or the Oceanidae as delimited in this publication. 
Forbes's usage was adopted by Kölliker (1853) when he described Oceania armata, 
and by Gegenbaur (1856) in his taxonomic system of the medusae. Agassiz (1862) 
returned to the usage of Lesson (1843) and with some ambiguity he even designated a 
type species. He noted for a conglomerate of species nowadays allocated to several 
families (Agassiz, 1862: 346, footnote 2): "I see, however, no reason why the name 
Nucleiferae, which he [Lesson] proposed for the old Forskalian type, should not be 
retained for this family, and the name Oceania and Oceanidae applied specifically, as 
Lesson has done, to Oceania phosphorica, which Péron & LeSueur place in the first 
section of the genus. This corresponds to the genus Thaumantias of modern writers." 
The sentence is somewhat ambiguous and it is debatable whether this must be seen as 



OCEANIDAE AND PACHYCORDYLIDAE 333 

a valid type-species designation (comp. ICZN § 67.5.3), but in the following taxo- 
nomic overview, Agassiz (1862) used the genus Oceania approximately in the sense of 
the present-day usage of Clytia. Oceania phosphorìca Péron & Lesueur, 1810, the 
species which Agassiz considered typical for the genus, is likely a synonym of Clytia 
hemisphaerica (see Goy, 1995). Following authors adopted either Forbes's or Lesson's 
view of the genus. This caused so much confusion that Haeckel (1879) abandoned the 
genus Oceania. In order to settle the dispute, and in order to stabilize the prevailing 
usage, Mayer (1910: 147) designated Oceania armata Kölliker, 1853 as type species 
for the genus Oceania. This settled the dispute and all following authors, notably also 
Kramp (1959, 1961, 1968) in his influential synopses, used Oceania as proposed by 
Mayer (1910). However, Mayer's type species designation is not in accordance with 
provisions of the ICZN currently in use (Calder, 1988b). The designation is invalid 
because O. armata was not one of the originally included species (ICZN § 69.1). This 
contrasts with the usage of the genus Oceania which was employed almost exclusively 
for the species Oceania armata, a circumglobal species that is well known and has 
regularly been reported in the literature of the last 150 years. Nomenclatural stability 
is thus severely threatened if the genus Oceania were to be replaced by a new name, 
or even worse, if recognized as a senior synonym of the leptomedusan genus Clytia 
Lamouroux, 1812. Therefore, after the publication of this study, a case will be made to 
conserve Oceania in its current meaning by asking the International Commission on 
Zoological Nomenclature to rule that Oceania armata is the type species of the genus, 
notwithstanding the fact that it was not an originally included species and overruling 
the potential type-designation of Agassiz. 

If the genus Oceania Péron & Lesueur, 1810 is accepted as valid, then auto- 
matically also the family group name Oceanidae Eschscholtz, 1829 becomes available. 

Turritopsis and Oceania medusae resemble each other quite closely, but they 
can be distinguished (see remarks under genus Turritopsis). 

Oceania armata Kölliker, 1853 Fig. 5 

Oceania armata Kölliker, in Gegenbaur et ah, 1853: 323; Metschnikoff, 1886: 78, pl. 1 figs 32- 
39; Mayer, 1910: 147, figs 80-81; Uchida 1927: 219; Kramp, 1959: 99, fig. 63; Kramp 
1961: 65; Kramp, 1968: 27, fig. 67; Brinckmann-Voss, 1970: pl. 4 flg. 2; Bouillon, 1980 
308; Bleeker & Van der Spoel, 1988: 230, flg. 6; Bouillon, 1995: 224; Schuchert, 1996 
15, fig. 4a-c. 

Oceania flavidula - Gegenbaur, 1856: 223, pl. 7: fig. 4. 

[not Oceania flavidula Péron & Lesueur, 1810 (= Laodice undulata (Forbes & Godsir, 1851))]. 

Turritopsis armata - Haeckel, 1879: 65. 

Callitiara polyophtalma Haeckel, 1879: 67, pl. 3 figs 1-5. 

Conis cyclophthalma Haeckel, 1879: 55, pl. 4 fig. 1; Hartlaub, 1913: 342, synonym; Kramp, 
1955: 152, synonym. 

Tiarella parthenopia Trinci, 1906: 208, figs. 

Material examined - MHNG INVE 32787, Naples, coll. 1892 - MHNG INVE 34177. Naples, 
det. Hartlaub, 1 medusa - MHNG INVE 31178, Villefrance-sur-Mer, France, det. 
Hartlaub, 5 medusae. 

Type material - There presumably exists no type material of O. armata. 

Description - Mature medusa 5-10 mm high, maximal bell diameter about the 
same as height, umbrella bell-shaped with flat top, jelly uniformly thin. Manubrium on 



334 



P. SCHUCHERT 




Fig. 5 
Oceania armata Kölliker, 1853, A-C, after preserved material from Naples; D, modified after 
Metschnikoff (1886). A) Mature medusa, scale equals 2 mm. B) Manubrium with parts of radial 
canals, scale equals 1 mm, g = gonad, mo = mouth, re - radial canal, rr = radial rib. C) Higher 
magnification of mouth rim fringed with pedicellate nematocyst clusters, scale 0.1 mm. D) 
Young hydranth, no scale available. 



OCEANIDAE AND PACHYCORDYLIDAE 335 

a transparent, shallow peduncle without vacuolated cells. Manubrium large, up to 2/3 
of subumbrellar height, ovoid, manubrium base constricted, with funnel-shaped mouth 
region. Stomach and mouth region distinctly cruciform in cross-section. Radial canals 
where joining manubrium funnel-like dilated and composed of large cells, continued 
on manubrium as radial ribs like four clasping claws (Fig. 5B). Mouth rim crenulated, 
with four prominent perradial lips. Margin of mouth with a continuous row of spherical 
nematocyst clusters, these usually on short pedicels. Gonads smooth, on interradial sur- 
face of stomach. Four quite broad radial canals, margins occasionally jagged, circular 
canal broad. Marginal tentacles tapering, 80- 1 20 in number, longer than bell (preserved 
sample), gastrodermis chordoid. Origins of tentacles somewhat alternately displaced 
adaxially and abaxially. Each tentacle with a slight proximal swelling beginning short- 
ly after origin. Adaxial ocellus near tentacle base. Nematocysts: microbasic euryteles, 
desmonemes. Colours (after Haeckel, 1879): Manubrium and gonads yellow or brown- 
yellow, mouth-lips orange or red, radial canals, ring canal and tentacles yellow, ocelli 
brown-red. Egg size about 0.27 mm. Polyp stage (after Metschnikoff, 1886) colonial, 
with creeping, ramified stolons. Hydranths on short, periderm-covered pedicel. 
Hydranth spindle-shaped with conical hypostome. With around 13 filiform tentacles in 
up to four whorls. 

Biology - Usually occurring in depths of less than 200 m, but not at surface 
(Kramp, 1965). 

Distribution - Mediterranean, coasts of Senegal and Gambia, Canary Islands, 
Cape Verde, Azores, Portugal, Spain; West Indies; Japan, New Zealand, Tasman Sea. 
The northern limit in Europe seems to be Portugal. Type locality: Mediterranean. 

Remarks - The polyp Oceania armata is known only from the rearing 
experiments of Metschnikoff (1886) and has never been found in nature. The origin of 
medusae buds is also not known. 

Bleeker & van der Spoel (1988) described the morphologically similar Oceania 
tydemani based on a single medusa taken south-west of the Azores. Oceania tydemani 
differs from O. armata in having more tentacles (180) and the nematocyst clusters of 
mouth margin are absent over four short interradial stretches. 

The identity of Oceania flavidula Péron & Lesueur, 1810 is pertinent to the 
question of the validitity of the genus Oceania (see remarks under genus Oceania). 
Gegenbaur (1856) believed O. flavidula and O. armata to be conspecific. The plates of 
Péron & Lesueur published by Goy (1995) now allow to identify O. flavidula as a 
synonym of Laodice undulata (Forbes & Godsir, 1851). Thus, the genus Oceania sensu 
Péron & Lesueur, 1810 did not originally include Oceania armata Kölliker, 1853. 

Corydendrium van Beneden, 1844 (emended) 

Synonym - Soleniopsis Ritchie, 1907. 

Type species - Sertularia parasitica Linnaeus, 1767. 

Diagnosis - Erect, branching or stolonal hydroid colonies, monosiphonic or 
polysiphonic. In branching colonies, branches and hydranth pedicels adnate to other 
branches over part or almost all their length. Perisarc firm, terminating near hydranth 
base, tubes of sister-branches nested. Hydranths spindle- to club-shaped; filiform 



336 P. SCHUCHERT 

tentacles scattered over much of hydranth body. Gonads formed within perisarc tubes 
of branchlets or at the end of branchlets and thus replacing a hydranth. Gonads gastro- 
dermal. 

Remarks - The taxononomic history of this genus was reviewed by Calder 
(1988a). Kramp (1935) has not been followed in merging Corydendrium with Turrl- 
topsis (see discussion above). As mentioned in the introduction to the family, the genus 
Corydendrium is here restricted to Oceanidae which have at least one of the following 
synapomorphies: (I) gonads within the perisarc tubes of hydranth pedicels or 
branchlets, (II) nested perisarc tubes. 

Clearly also a member of the genus Corydendrium in the sense of the author is 
the Japanese C. brevicaulis Hirohito, 1988. This species is quite important in dis- 
cussions of the scope of the genus as it forms either stolonal or branched colonies. In 
its morphology, this species otherwise resembles C. parasiticum so closely that it could 
arguably seen only as a variant of it. The generic diagnosis was therefore modified to 
include stolonal colonies to permit the inclusion of C. brevicaulis Hirohito, 1988. 

Hirohito (1988) also described another Corydendrium, namely C. album. It 
resembles closely Turritop soldes brehmeri Calder, 1988b and should be transferred to 
the genus Turritop soldes if the so far unknown female gonophores of C. album will 
prove to be identical. 

Fiorini (1977) argued that the reproductive organs of C. parasiticum are pro- 
bably not homologous to a gonophore and thus to a vestigial medusa. They are most 
likely a composite structure derived from a polyp with several gonophores (the term 
gonophore must be restricted to gonad-bearing structures that can be homologized with 
a single medusa bud). Such a composite gonad-bearing organ is thus better referred to 
as a gonangium (see also discussion in Thiel, 1962). Contrary to most other hydro- 
zoans, the germ cells of Corydendrium parasiticum and of the males of C. dispar 
remain in the gastrodermis (Weismann, 1883; Fiorini, 1977; Kramp, 1935). Other 
species referred here to Corydendrium should be examined if they have similar 
gonangia, which would consolidate it as an additional synapomorphy for the genus. 
Germ cells located in the epidermis represent a synapomorphy for the Hydrozoa 
(Schuchert, 1993). The gastrodermal gonads of some Corydendrium species must be 
seen a character reversal. All other related taxa have epidermal gonads. 

Corydendrium corrugatimi Nutting, 1905 from Hawaii likely also belongs to 
the genus Corydendrium, although its reproductive structures remain unknown (see 
Schuchert, 2003b). The same holds true for C. flabellatum Fraser, 1938 from the 
tropical eastern Pacific and C.fruttcosum Fraser, 1914 from Vancouver Island (north- 
eastern Pacific). Corydendrium splendidum Boone, 1938 is a synonym of Pennaria 
dlstlcha Goldfuss. 1820 (see Cooke, 1977). 

The European fauna includes only two species, namely Corydendrium para- 
siticum (Linnaeus, 1767) and Corydendrium dispar Kramp, 1935. 

Corydendrium parasiticum (Linnaeus, 1767) Fig. 6 

Sertularia parasitica Linnaeus. 1767: 1315. 

? Solenopsis dendriformis Ritchie, 1907: 495, figs 142-143, pi. 26 fig. 1. 

? Corydendrium sessile Ritchie, 1910a: 802, pi. 76 figs 1-2. 



OCEANIDAE AND PACHYCORDYLIDAE 



337 




Fig. 6 
Corydendrium parasiticum (Linnaeus, 1767), after preserved material from Naples. A) Colony 
silhouette, scale equals 1 cm. B) Distal branches with two hydranths, note gonads (stippled) in 
pensare tube of pedicels and adnate braches, scale equals 1 mm. 



not Clava (Corydendrium) parasiticum - Bonnevie, 1899: 39 [= Corydendrium dispar Kramp, 
1935]. 

Corydendrium parasiticum - Weismann, 1883: 34, fig. 7, pi. 14 figs 1-9, pi. 15 figs 1-5, pi. 16 
figs 1-2, 4; Neppi, 1917: 43, figs 7-8; Rossi, 1971: 20, fig. 6L; Millard, 1975: 72, fig. 
24B-D; Wedler & Larson, 1986: 82, fig. 5C; Rees & Vervoort, 1987: 12, bibliography: 
Calder, 1988a: 6, figs 3-4; Hirohito, 1988: 66, fig. 21 a-c, pi. 2 fig. B; Migotto, 1996: 11, 
fig. 2e; Watson, 1999: 2, fig. 2A-E; Calder et al. 2003: 1176, fig. 2. 

Material examined - MHNG INVE 25127, Naples, Italy, fertile males, very profuse colony - 
MHNG INVE 31489, Naples, Italy, coll. 1892 - MHNG INVE 31490, Naples, Italy, coll. 
1896. 

Type material - No type material was localized. 

Description - Colonies bushy, much branched, polysiphonic, 2-5 cm in height 
(reportedly up to 16 cm). Perisarc stiff, smooth or encrusted with silt, not annulated, 
terminating below hydranths, sometimes terminal part slightly dilated and wrinkled. 
Branches adnate for some distance, then diverging at acute angle. Perisarc tubes of 
side-branches nested at their origins. Diameter of branches 0.4-0.5 mm, terminal tubes 
0.3 mm. Hydranths elongated, cylindrical to fusiform, 1-3 mm long, diameter up to 
0.35 mm, with 20-40 filiform tentacles scattered over most of hydranth body. 



338 P- SCHUCHERT 

Hypostome prominent, dome-shaped. Sexual organs tubular structures, length 3 mm 
and more, arising as side branches of hydranth pedicels, initially wholly contained 
within perisarc tube, mature eggs extruded from perisarc but remaining attached near 
end of tube until developed into planulae, thus larviparous (Neppi, 1917; Wedler & 
Larson, 1986). Structure of sexual organs simple, bi-layered, gametes in gastrodermis. 
Nematocysts: microbasic euryteles and desmonemes. 

Biology - Grows most often on rock. The fertile period in the Mediterranean 
is from August to September, while colonies are present from June to October (Lo 
Bianco, 1909). Motz-Kossowska (1905) obtained fertile colonies from the Balearic 
Archipelago in August. Neppi (1921) found colonies during June and August, and they 
were also fertile during this period. In the Mediterranean, the species seems to be only 
locally abundant, with most records coming from Naples. Intensive searches for it at 
localities near Banyuls-sur-Mer (Motz-Kossowska, 1905), Island of Medes (Gili, 
1982) and near Genoa (Boero & Fresi, 1986) were unseccussful. 

Other information - The microscopic structure was investigated in detail by 
Weismann (1883) and Fiorini (1977). The ultrastructure of the planula and the oocytes 
were investigated by Glätzer (1970, 1971). The early development was examined by 
Neppi (1917) and Glätzer (1971). 

Distribution - Perhaps circumglobal in tropical and subtropical seas: along the 
European coasts confined to the Mediterranean (Weismann, 1883; Neppi, 1917), also 
coasts of western and southern Africa (Ritchie, 1907; Millard, 1975), tropical western 
Atlantic (Wedler & Larson, 1986; Calder, 1988a; Migotto, 1996), Indian Ocean 
(Millard & Bouillon, 1973; Rees & Vervoort, 1987), Indonesia (Vervoort, 1941; 
Schüchtert, 2003b), coasts of northern Australia (Watson, 1999), tropical western 
Pacific (Leloup, 1937, as C. dendriforme), north-western Pacific (Hirohito, 1988), ? 
tropical eastern Pacific (Fraser, 1938, as C.flabellatum). Depth range about 1-90 m. 

Remarks - The taxonomic history and synonymy of this species are given by 
Calder (1988a) and need not be repeated here in detail. Vervoort (1941), Millard 
(1975), and Calder (1988a) considered Corydendrium dendriforme (Ritchie, 1907) 
from the Cape Verde Islands conspecific with C. parasiticum. This contrasts with Rees 
& Thursfield (1965) who re-examined the type material of C. dendriforme and kept it 
distinct. Both species reportedly differ in the more arborescent colony of C. dendri- 
forme versus the more bushy one of C. parasiticum (see fig. 6). Although the argu- 
ments in favour of a conspecificity appear more convincing, a definite answer is not 
possible at the moment. A detailed comparison of the populations from the Cape Verde 
Archipelago with those from adjacent regions is needed to provide the necessary argu- 
ments. Corydendrium dendriforme (and also C. sessile) are thus treated as questionable 
synonyms of C. parasiticum. The material from the Strait of Gibraltar reported by 
Ramil & Vervoort (1992) as C. parasiticum is somewhat atypical and perhaps does not 
belong to this species. 

Corydendrium dispar Ksamp , 1935 Fig. 7 

Clava (Corydendrium) parasiticum - Bonnevie, 1899: 39. 
[not Corydendrium parasiticum (Linnaeus, 1767)]. 



OCEANIDAE AND PACHYCORDYLIDAE 339 

Corydendrium dispar Kramp, 1935: 1, figs 1-4. 

not Corydendrium dispar - Rasmussen, 1973: 22 [ = Pachycordyle navis (Millard, 1959)]. 

Type Material examined - ZMUC, holotype, Kungsbackafjord, Sweden, coll. 26 Jun 1928, 26 
m - ZMUC, paratype, Kvittings0y, Norway. 

Other Material examined - BIOFAR collection (Norrevang et a/., 1994), kept by Kaldbak 
Marine laboratory, The Faroes: Station 8 (62.29°N 5.40°E, 171 m, 17 Jul 1987, on bi- 
valve shell); station 165 (62.18 °N 4.97°E, 184 m, 07 May 1988), slides of this material 
deposited as MHNG LNVE 33555; station 204 (62.16°N 5.88°E, 158 m, 11 May 1988, 
on Abietinaria fiisca). 

Description - (after Kramp, 1935 and examined material) Colonies forming 
delicate, erect shoots, less profuse than C. parasiticum, 0.5 - 2 cm in height, reportedly 
up to 6.5 cm. Stolons creeping, ramified, anastomosing. Shoots irregularly branched, 
entirely monosiphonic or polysiphonic in lower parts of the shoots, branches adnate to 
each other over variable distances, diameter of single branch up to 0.2 mm, perisarc 
tubes at origin of branches nested, perisarc wrinkled, generally double-layered, inner 
layer thick and firm, outer layer membranous. Hydranths spindle-shaped, 0.6 mm in 
preserved material, hypostome dome-shaped, 16-20 filiform tentacles scattered over 
most of hydranth body. Reproductive organs not within perisarc tubes of branchlets, 
arising like hydranths at end of side-branches, covered by very thin perisarc mem- 
brane. Male sexual organs are sessile sporosacs, oblong-elliptic or -ovoid, length 0.7 
mm, diameter 0.4 mm, on short pedicels; structure sac-like, with thin epidermis and 
thick gastrodermis containing the gametes. Mature female organs insufficiently 
known, likely sessile sporosacs, spherical, diameter up to 1.2 mm. 

Biology - Known substrates are shells and hydroids (Kramp, 1935; this study). 
Occurs in fully marine environments only. Depth range: 26 to 184 m. 

Distribution - Along Swedish and Norwegian coast north to Trondheim Fjord 
(Kramp, 1935), The Faroes. Type locality: Kungsbackafjord, south of Gothenburg, 
Sweden, 26 m (Kramp, 1935). 

Remarks - Corydendrium dispar Kramp, 1935 is recorded here for the first 
time in waters of the Faroe Islands. The samples contained only male specimens and 
the mature female reproductive structures are still imperfectly known. While the male 
structures are only simple sacs without any recognizable vestiges of a medusa 
structure, the initial developmental stages of female ones suggest a medusoid origin 
(Kramp, 1935). 

The material recorded by Rasmussen (1973) as C. dispar was re-examined and 
identified as P. navis. See remarks under this species. 

Merona Norman, 1 865 

Type species - Tubiclava cornucopiae Norman, 1864 by monotypy. 

Diagnosis - Stolonal hydroid colonies, polyps polymorphic with gastrozooids. 
gonozooids and nematophores. Hydranths with scattered filiform tentacles and 
perisarc-covered caulus. Gonozooids reduced, without mouth or tentacles, thus blasto- 
style-like, bearing numerous gonophores. Gonophores sessile sporosacs. Defensive 
zooids small, arising from stolons, in funnel-shaped perisarc tube. 



340 



P. SCHUCHERT 




Fig. 7 
Corydendrium dispar Kramp, 1935; A, after paratype material; B-C, after preserved material 
from The Faroes. A) Part of shoot, the putative female sporosacs were empty in this material, 
scale equals 0.5 mm. B) Branch with young male reproductive structure, optical section, scale 
bar 0.2 mm. C) Mature male reproductive organ, optical section, same scale as B. 

Remarks - The gonophores of Merona arise on blastostyles that can either be 
interpreted as being derived evolutionary from a hydranth or from a hydranth pedicel. 
Their simple structure does not lend itself to an easy answer. If the blastostyles are de- 
rived from a hydranth body, this would be problematic because in all other Oceanidae 
the gonophores arise on the pedicels or stolons (see above). The alternative interpreta- 
tion that the gonozooids are homologous to hydranth pedicels only is much better com- 
patible with the scope of the family as outlined in this publication. 



Merona cornucopiae (Norman, 1864) 



Fig. 8 



Tubiclava cornucopiae Norman, 1864: 82, pi. 9 figs 4-5; Hincks, 1868: 11, pi. 2 fig. 2; Allman, 

1872: 258; Fraser, 1937: 23, fig. 4; Fraser, 1944: 36, fig. 4. 
Merona cornucopiae - Broch, 1916: 40, pi. 2 fig. 17, fig. L; Rees, 1956: 499, figs 1-3; Cabioch, 

1965: 401, figs 1-3; Millard & Bouillon, 1973: 28, fig. 3H-J; Millard, 1975: 74, fig. 25; 

Medel. Garcia-Gómez & Bouillon, 1993: 515. figs 1-2B; Ramil et al., 1998: 184, fig. 

2a-b. 

Material examined - ZMUC, Denmark, Groves Flak, 25 m, 17 May 1923, on scaphopod. det. 
Kramp, infertile - ZMUC, Faroe Islands, Skalbund, 7 Jun 1899, on bivalve shell, much 



OCEANIDAE AND PACHYCORDYLIDAE 



341 






Fig. 8 
Merona cornucopiae (Norman, 1864), after preserved material from Roscoff. A) Hydranth, quite 
expanded, scale equals 0.2 mm. B) Nematothecae on stolons, same scale as A. C) Blastostyle 
with female gonophores, scale equals 0.2 mm. D) Female gonophore, scale equals 0. 1 mm. 



damaged, with blastostyles - IRSNB, France, Brittany, Roscoff, numerous colonies on 
shells of Turritella spec, and Nassarius spec, inhabited by Phascolion spec, not well 
preserved, one blastostyle seen - BMNH 1956.11.7.1, Brittany, Baie de Morlaix, off. 
Callot, 22-23 Jun 1955, 15-25 m, on bivalve and Turritella, coll. W. J. Rees (material of 
Rees, 1956), one colony with female gonophores - BMNH 1941.3.20.516, English 
Channel, Eddystone, 4 Jun 1898, coll. E. T. Browne (material described in Rees, 1956), 
soft tissues quite damaged, several blastostyles with female gonophores present. 
Type material - The type material of M. cornucopiae is kept by the BMNH (Rees, 1956). 

Description - (after examined material, and Rees, 1956) Colonies stolonal, 
never branched, stolons creeping, reticulated or coalesced into mat. Gastrozooids 
consisting of a perisarc-covered pedicel and a terminal hydranth. Living tissue of 
hydranth pedicels not well demarcated from hydranth body. Pedicels in wide perisarc 
tube, the latter 1-5 mm in height, diameter at distal end double or more the basal 
diameter (0.1-0.14 mm), perisarc thus cone-shaped, often slightly bent or curved, 
irregularly corrugated, often coated by mud and fine sand grains. Contracted hydranth 
able to withdraw almost completely into perisarc tube. Relaxed hydranths 1-5 mm. 



342 P. SCHUCHERT 

rod- to slightly spindle shaped, with 16-20 long, filiform, tapering tentacles scattered 
over distal half or more. Hypostome dome-shaped. High gastrodermal cells in 
proximal half of hydranth body, apparently vacuolated. Colonies dioecious. Gono- 
phores arise on blastostyles with a short basal perisarc cone as in the hydranths. 
Gonophores in a dense cluster at distal end of blastostyle. In females, blastostyles with 
pointed tip that may slightly overtop gonophore cluster, 15-20 gonophores per 
blastostyle. Gonophore ovoid when mature, length up to 0.2 mm, with distinct stalk, 
continued through gonophore as spadix, distal part of spadix slightly protruding like 
nipple, up to six eggs per gonophore. Male blastostyles (after Medel et al., 1993) 
shorter, gonophores more dense, spherical, spadix perhaps (?) not protruding. Nema- 
tophores facultatively present, arising from stolons, about 0.3 mm in height, simple 
tentacle-like, within thin perisarc tube, the latter at distal end widening funnel-like, tip 
of nematophore with concentration of large euryteles. Nematocysts (after Cabioch, 
1965): large microbasic eurytele (19-22) x (5) pirn, discharged shaft about two times as 
long as capsule; medium sized microbasic eurytele (13-14) x (5) //m, shaft as long as 
capsule, mainly on tentacles; small microbasic eurytele, (8) x (3) pm, shaft as long as 
capsule, mainly on tentacles; desmonemes (5) x (3) pm. Colours in life: Hypostome 
intense white, gastrodermis of hydranth body-orange brown to scarlet, fading in non- 
tentacular portion into a uniformly pale tint. 

Biology - Merona cornucopiae grows predominantly on shells of molluscs 
such as Glycimeris, Aporrhais, Dentalium, Turritella and others, very often on Turri- 
tella shells inhabited by the sipunculid Phascolion strombi. Occurrence is on sediment 
bottoms in depths of 10-274 m. Fertile specimens were collected at Plymouth in May 
and June (Rees, 1956), at Roscoff June-July (Teissier, 1965), at Oslofjord end of April 
(Christiansen, 1972), at Trieste January-February (Graeffe, 1884). 

Distribution - British Isles (Rees, 1956), Brittany (Rees, 1956; Cabioch, 
1965), Faroes (Broch, 1916), Norway (Christiansen, 1972), Spain (Ramil et al, 1998; 
Medel & Lopez Gonzalez, 1996), Mediterranean (Graeffe, 1884), north-western 
Atlantic (Fraser, 1944), north-eastern Pacific (Fraser, 1937), South Africa (Millard, 
1975), Seychelles (Millard & Bouillon, 1973). Type locality: Shetland Islands. 

Remarks - Medel et al. (1993) described what they considered a sexual 
dimorphism of the blastostyles of Merona cornucopiae. They found that the female 
blastostyles are longer and they have their gonophores distributed over a larger area, 
resulting in a less dense arrangement than in males. In the present study, only females 
were observed and the findings of Medel et al. could therefore not be verified. It 
nevertheless appears that some of the differences are probably only due to different 
states of contraction and different ages. Rees (1956: figures 1 and 2) shows both types 
of blastostyles, but both are females (a re-examination of Rees's material confirmed 
this). The apparent density of the female sporosacs also depends also on how many 
eggs there are present. A partial spawing results in an apparent reduction of the density 
of sporosacs, because only the thin spindle-shaped spadices remain of the initially 
oviform sporosacs. 

Merona ibera Medel, Garcia-Gomez & Bouillon, 1993 and M. cornucopiae can 
be distinguished only by their female gonophores. At least for the Mediterranean, it is 



OCEANIDAE AND PACHYCORDYLIDAE 343 

therefore not possible to reliably identify Merona species without female gonophores. 
Some of the records of Merona cornucopiae from the Mediterranean (e. g. Motz- 
Kossowska, 1905) are thus perhaps referable to M. ibera. Graeffe's (1884, as Tubiclava 
cornucopiae) material from Trieste had four eggs per gonophore. It thus corresponds 
more with M. cornucopiae than with M. ibera. 

Merona ibera Medel, Garcia-Gomez & Bouillon, 1993 

Merona ibera Medel et al, 1993: 513, figs 2C-D & 3A-C. 

Type material - The type material of M. ibera is kept by the Museo Nacional de Historia 
Natural (Madrid) and the RMNH. 

Diagnosis - Similar to Merona cornucopiae, but female blastostyles smaller 
and gonophores placed more distally, eggs smaller (37-60 /mi), more eggs per 
gonophore (>12 per gonophore), female gonophore spindle-shaped, distal end of 
spadix with button of nematocysts. 

Distribution - Known from type locality only: Algecirras Bay, Strait of 
Gibraltar, Mediterranean, 21 m, on bivalve Gouldia minima. 

Remarks - See under M. cornucopiae. 

Rhizogeton L. Agassiz, 1 862 

Type species - Rhizogeton fusiformis L. Agassiz, 1862. 

Diagnosis - Stolonal hydroid colonies, polyps monomorphic, hydranths either 
sessile or with a perisarc-covered caulus. Hydranths with scattered filiform tentacles 
covering at least half of hydranth body. Nematophores absent. Gonophores sessile 
sporosacs arising from stolons. 

Remarks - Calder (1988a) thought that it might be necessary to split this genus 
and proposed the new genus Rhizodendrium. He held the genus Rhizodendrium distinct 
from Rhizogeton on account of the following characters: the hypostome is short and 
dome-shaped instead of very elongate and conical, it has 20 or more tentacles instead 
of about 10, its gonophores are more rounded than fusiform. The gonophores of 
Rhizodendrium sterreri Calder, 1988, the type species of the genus Rhizodendrium, are 
so far unknown. The difference in the shape of the gonophores was therefore based on 
Rhizogeton nudus and R. eozense Yamada, 1964, which according to Calder (1988a) 
should be transferred to the genus Rhizodendrium. Although the differences in the 
shape of the gonophore are not so apparent (Dons, 1912), Rhizogeton fusiformis 
appears otherwise to be distinct from R. nudus (comp. Fig. 9 and figure 3 in Fraser, 
1944). The proposal of Calder (1988a) to separate them into two genera could there- 
fore be justified. If the gonophores of R. sterreri are discovered and independent data 
including DNA sequences show that Rhizogeton and Rhizodendrium species do not 
form a monophyletic group, then Calder's proposal should be followed. But because 
the gonophores of the type species of Rhizodendrium are so far unknown, and in order 
to maintain nomenclatural stability, only the genus Rhizogeton is used here. 

Rhizogeton nudus Broch, 1909 Fig. 9 

Rhizogeton nudum Broch, 1909: 137, fig. 1; Antsulevich & Polteva, 1986: 967, fig. 2; Jones, 
1992: 721, fig. 1; Schuchert, 2001a: 20, fig. 10A-C. 



344 P. SCHUCHERT 

Rhizogeton fusiformis - Browne, 1897: 243. 

[not Rhizogeton fusiformis L. Agassiz, 1862]. 

Rhizogeton nudus - Dons, 1912: 53, fig. A-C; Calder, 1972: 223, pi. 2 fig. 1. 

Rhizodendrium nudum - Calder, 1988a: 11. 

Material examined - Material from Greenland and Iceland, see Schuchert (2001a). 
Type material - The type material of R. nudus could not be located. 

Description - Colonies stolonal, stolons creeping, ramified. Hydranths appear- 
ing sessile, without distinct pedicel, at base a short beaker of very thin perisarc, usually 
invisible. Hydranths up to 2-5 mm in height, rod-shaped, with 14-19 tentacles, these 
scattered in distal half to distal third of hydranth. Tentacles with chordoid gastrodermis, 
tentacles smooth, tapering, quite long and thin, lengths unequal, proximal ones shorter. 
Hypostome dome-shaped. High gastrodermal cells in proximal half of hydranth body, 
apparently vacuolated. Gonophores arising from stolons and developing into sessile 
sporosacs without canal system or velum but with spadix; with short pedicel enveloped 
by perisarc, young sporosacs also enveloped by very thin, filmy perisarc. Female 
sporosacs ovoid to spindle-shaped, diameter 0.35-0.45 mm, length 0.6-0.7 mm, with 3- 
5 eggs, at maturity the sporosac envelope disappears but eggs and developing embryos 
stick to the spadix. Male sporosacs oblong ellipsoids, about the same size as female 
sporosacs. Nematocysts: microbasic euryteles, 7x3 //m; desmonemes 5x3 pim. 
Colour in life: pinkish. 

Variation - Females sporosacs from Britain may contain up to 15 eggs, 
hydranths are orange (Jones, 1992). According to Dons (1912) the male sporosacs 
become ovoid with a pointed distal end when fully mature. 

Biology - Grows on a variety of substrates such as mussels, hydroids, 
bryozoans, and holdfasts of macroalgae. Depth range 0-100 m. 

Distribution - Greenland (Kramp, 1914; Schuchert, 2001a), Iceland 
(Schuchert, 2001a), Spitsbergen (Broch, 1909), White Sea (Antsulevich & Polteva, 
1986), Norway (Dons, 1912), British Isles (Jones, 1992), north-eastern Canada 
(Calder, 1972). Type locality: Bismarck Sound, Spitsbergen, 35 m, on Grammaria 
abietina. 

Remarks - In his first description, Broch (1909) used the specific name nudum. 
In a footnote, Dons (1912) mentioned that Broch had asked him through a letter to 
change this to nudus because the gender of Rhizogeton is masculine ("geton" is derived 
from the Greek word yenwv for neighbour, which is masculine). Following the rules 
of the ICZN (ICZN, 1999: § 31.2 and 34.2), the specific epithet must have an ending 
matching the gender of the genus. The correct name is thus Rhizogeton nudus Broch, 
1909. 

Broch (1909) and Dons (1912) described the hydranths as devoid of a basal 
perisarc. Careful examination with the help of a compound microscope nevertheless 
revealed the presence of a fine perisarc in the material examined here. 

The development of the sporosacs is well documented by the study of Dons 
(1912). 

The sporosacs depicted by Millard (1975) had no pedicels and were directly 
attached to the stolons. This material from the tropical Indian Ocean as well as the 



OCEANIDAE AND PACHYCORDYLIDAE 



345 



PSt 




Fig. 9 
Rhizogeton nudus Broch, 1909, after preserved material from Iceland (A) and Greenland (B-C). 
A) Hydranth and presumed gonophore, scale equals 0.3 mm. B) Female sporosacs, right with 
shed envelope and eggs still sticking to spadix, scale equals 0.5 mm. C) Young male sporosac, 
same scale as B. 



records of Mammen (1963) and Ritchie (1910b) may therefore belong to separate 
species. Mammen (1963), Rees & Thursfield (1965) and Antsulevich & Polteva (1986) 
made similar suggestions. The male sporosacs described by Genzano (1993) in 
material from Argentina likewise lacked a pedicel and may therefore also belong to 
another species. Rhizogeton nudus is probably restricted to arctic and cool-temperate 
waters of the North Atlantic. 

Park (1997) suggested that R. fusiformis L. Agassiz, 1862 and R. nudus are 
conspecific. Although Dons (1912) kept both species apart, he found that there is little 
difference in the morphology of the gonophores. Calder (1988a) re-examined the type 
material of R. fusiformis and he confirmed that this species has a pointed hypostome 
and only about 10 tentacles. The differences even prompted Calder (1988a) to place the 
two species in different genera (see remarks under genus Rhizogeton). I therefore think 
that R. fusiformis and R. nudus of the north-west Atlantic should be kept distinct. 

In his report on hydroids of Valencia Harbour (western Ireland), Browne (1897) 
also described material which he attributed to R. fusiformis L. Agassiz, 1862, although 
he noted marked differences to Agassiz's description. The most critical difference was 



346 P. SCHUCHERT 

the number of tentacles, which was around 20 and not 10-12. The data given by 
Browne (1897) closely match R. nudus as described above and it is very likely that his 
specimens belonged to this species. This interpretation is supported by the finding of 
R. nudus in the Morecambe Bay (Irish Sea, England; Jones, 1992). 

Cordylophora Allman, 1 844 

Type species - Cordylophora lacustris Allman, 1844 by monotypy. 

Diagnosis - Erect, branching hydroid colonies, side-branches and hydranth 
pedicels not adnate, pensare tubes not nested. Relaxed hydranths spindle-shaped, 
hypostome well delimited from hydranth body, filiform tentacles scattered in distal half 
of hydranth body. Gonophores oblong, borne on pedicels of hydranths and on 
branches, sessile sporosacs with branched spadix. 

Remarks - Cordylophora resembles somewhat to the genus Pachycordyle and 
could as well belong to the Pachycordylidae (comp. Moiri, 1980; Stepanjants et al., 
2000). Unfortunately, there are no convincing synapomorphies that would link it 
unambiguously to either the Oceanidae or the Pachycordylidae. Mainly for historic 
reasons, it is here kept in the Oceanidae. It shares with this family the tentacles that are 
scattered over much of the hydranth body, while in the Pachycordylidae they are 
concentrated in the distal part. 

Cordylophora caspia (Pallas, 1771) Fig. 10 

Tubularia caspia Pallas, 1771: 479. 

Tubularia cornea Agardh, 1816: 258. 

Cordylophora lacustris Allman, 1844: 330; Allman, 1853: 367, pis 25-26; Schulze, 1871: 1-52, 

pis 1-6; Hincks, 1868: 16. pi. 3 fig. 2; Allman, 1872: 252, pi. 3; Nutting, 1901: 327, fig. 

2; Roch, 1924a: 350, footnote 1. 
Cordylophora albicola Kirchenpauer, in Busk, 1861: 284, pi. 9 figs 12-14; Allman, 1872: 254. 
Cordylophora lacustris var. otagoensis Fyfe, 1929: 813, figs 2-10. 
Cordylophora americana Leidy, 1870: 113. 

Cordylophora whiteleggi von Lendenfeld, 1886: 97, pi. 6 figs 11-12. 
Cordylophora lacustris otagoensis - Ralph, 1953: 64 fig. 6. 
Cordylophora caspia - Vervoort, 1946: 1 19, fig. 47b & 48a (not others); Naumov, 1969: 196, fig. 

66; Rossi, 1971: 20, fig. 61; Morii, 1980: 155, figs 1-3; Moni, 1981: 45, fig. 13, pi. 1 fig. 

3, pi. 2 fig. 3; Morii & Boero, 1986: 34, figs 14-15a-b; Barnes, 1994: 60, fig; Holstein, 

1995: 95 figs 49-50; Schuchert, 1996: 15, fig. 3a-e. 

Material examined - MHNG INVE 34179, Rostock, Germany, Baltic Sea (not well preserved, 
no hydranths left) - ZMUC, Randers Fjord, Mellerup, Denmark, 1.5 m, 25.05.1916 - 
ZSM, as C. lacustris, Elbe estuary, Germany, fertile - ZSM, as C. lacustris, Brunsbiittel, 
Germany, fertile - ZSM, as C. lacustris, Woltersdorf near Berlin, Germany, fertile, 
3.08.1920 - ZSM, as C. lacustris, Heringsdorf, Germany, fertile - ZSM, as C. lacustris, 
Greifswald, Germany, June 1902 - New Zealand, see Schuchert (1996). 

Type material - There presumably exists no type material of C. caspia. 

Description - Erect, branching hydroid colonies, stems up to 3 cm high 
(reportedly up to 10 cm), arising from reticulate stolons. Occasional autoepizooism 
possible. Erect shoots with monopodial growth, branching several times irregularly at 
various angles, ending in up to 40 hydranths. Shoots often composed of a long main 
stem and shorter side branches, branches not adnate, stem diameter about 0.2 mm. 



OCEANIDAE AND PACHYCORDYLIDAE 



347 




Fig. 10 
Cordylophora caspia (Pallas, 1771), A samples from Germany, B-C from Denmark. A) Colony 
silhouettes demonstrating variability, scale equals 5 mm. B) Hydranth with two gonophores on 
its pedicel, scale equals 0.2 mm. C) Schematic optical section of a mature female gonophore, 
eggs are shown stippled and surrounded by solid line, note branched spadix, scale equals 
0.2 mm. 



Colonies living under unfavourable conditions remain stolonal with hydranths on a 
simple pensare covered pedicel. Perisarc smooth with occasional annulated stretches, 
perisarc ends below hydranths. Relaxed hydranths spindle-shaped, contracted ones 
ovoid, 1-2 mm high, size variable even within a single shoot, with prominent hypos- 
tome, the latter bullet-shaped to conical. Scattered filiform tentacles, usually around 
14-16 (up to 27 possible), proximal third of hydranth body free of tentacles. Tentacles 
of varying length (up to 1.4 mm), tapering slightly to 3/4 of basal diameter, gastro- 
dermis composed of chordoid cells. Colours: hydranths white or pale pink, perisarc 
brown. Nematocysts: microbasic euryteles and desmonemes. Colonies dioecious. 
Gonophores ellipsoid, arising at pedicels of hydranths, branches and stems, covered 
with perisarc, mature about 0.7-1 mm long, diameter about half the length. Initially 
with a branched spadix, reduced in later development, gametes released through distal 
hole. Females with 7-16 eggs (fewer also possible), 0.07-0.12 mm in diameter, 
developing to planulae in situ. 

Occurrence - Circumglobal in temperate and subtropical regions, usually in 
brackish waters as in estuaries, river deltas and lagoons. Absent from fully marine 
environments. Sometimes penetrating into completely fresh water, but there usually a 
reduced development. When occurring in inland rivers and lakes, these water bodies 



348 P. SCHUCHERT 

are generally characterized by elevated concentrations of some ions (Arndt, 1984). 
However, it has recently also been found that C. caspia can successfully invade pure 
freshwater biotopes (Smith et ah, 2002). Depth range 0-85 m (Naumov, 1969), usually 
in very low waters, on rock, under stones, on Dreissena spec. (Bivalvia). Type locality: 
Caspian Sea. 

Remarks on variation, morphology and ecology - There is a vast body of 
literature on this species and it is impossible to review it here completely. Only the 
most important works are therefore mentioned. The morphology was studied in detail 
by Allman (1853), Schulz (1871), and Weismann (1883). The embryology was 
examined by Van de Vyver (1967). Roch (1924a, 1924b) reviewed all records of this 
species and examined the influence of chemical and physical parameters on the 
morphology. These studies were later extended by Kinne (1956, 1957) and Arndt 
(1984). The morphology of the colony and the hydranths correlates strongly with 
salinity. Near fresh-water conditions causes low colonies and stolonization, low 
salinities produce spherical hydranths with short, stubby tentacles, while in water with 
high salinity hydranths are elongated and their tentacles long and thin. A salinity of 
16.7 %o at 20 ° and 10 %o at 10 °C was found to be optimal for growth in the population 
examined by Kinne (1957, North Sea to Baltic Sea Channel). Arndt (1984) found a 
salinity optimum of 6-10 %c for populations of the Baltic and Arabian Sea. Sexual 
reproduction started when water temperatures rose above 10 °C. The temperature 
optimum was 11-18 °C. Smith et al. (2002) found the species in fresh-water and 
demonstrated that it has successfully invaded this biotope by adjusting its physiology 
and ecological needs. Cordylophora caspia is able to survive long periods without 
food. This is especially pronounced in fresh water, where it has lived without feeding 
up to six months (Kinne, 1956). Bouillon (1963) found that this species can be either 
larviparous or viviparous. For more references see also Holstein (1995), Folino (2000), 
and Smith et al. (2002). 

Taxonomic remarks - Although the original description of Tubularia caspia 
by Pallas (1771) is too vague to allow an unambiguous recognition, most contemporary 
authors have followed Roch (1924b) in considering it a senior synonym of C. lacustris 
Allman, 1844. Stepanjants et al. (2000) disagree, however, without providing 
arguments. A good review, and remarks on the taxonomic status, can be found in Folino 
(2000), who also considers the possibility that more than one species might be included 
in the present concept of C. caspia. It is certainly possible that C. caspia and C. lacus- 
tris - the latter referring to populations confined to nearly or fully freshwater 
conditions - are morphologically inseparable, but nevertheless represent two separate 
biological species. Genetic methods must be used to clarify the status of the various 
Cordylophora ecomorphs. 

Vervoort (1964) examined type material of Bimeria (?) baltica Stechow, 1923 
and found it indistinguishable from C. caspia. 

Pachycordylidae Cockerell, 1911 
Synonym - Clavopsellidae Thiel, 1962. 

Diagnosis - Anthoathecata Filifera, polyp phase forming stolonal or branching 
colonies, hydranths pedicellate, perisarc terminating at base of hydranth, without 



OCEANIDAE AND PACHYCORDYLIDAE 349 

pseudohydrotheca. Hydranths club-shaped, spindle-shaped, or amphora-shaped; 
hypostome nipple- to dome-shaped. Tentacles in two or more whorls restricted to a 
narrow band below hypostome, spreading over less than one third of the hydranth body 
only. Gonophores arising on stems or pedicels. Gonophores free medusae, medusoids 
that can be liberated, or fixed sporosacs. Medusae at liberation with four simple or 
branched oral tentacles arising above mouth, four tentacle bulbs with one tentacle each, 
ocelli present. 

Remarks - The family Pachycordylidae comprises the genera Pachycordyle 
and Silhouetta. Thiel (1962) proposed the new family Clavopsellidae comprising the 
genera Clavopsella, a synonym of Pachycordyle, and Balella Stechow, 1919. The latter 
genus has hydranths with two widely separated whorls of tentacles and it produces 
gonophores on gonozooids. Schuchert (2003b) argued that Balella has closer affinities 
with the Hydractiniidae and must be placed into a family of its own (Tubidendridae, 
although the valid name is Balellidae, see Calder 1988a: 13). Thiel was probably 
unaware that there was already a family group name available that included Pachy- 
cordyle, namely Pachycordylini Cockerell, 1911. Calder (1988a) raised it to subfamily 
level as Pachycordylinae and he provided arguments why it must be preferred over 
Clavopsellidae and why Silhouetta should be included in the group. Here, the taxon 
Pachycordylinae is raised to family level, becoming thus Pachycordylidae. I think this 
is necessary because Pachycordyle is so intermediate between Oceanidae and Bougain- 
villiidae that it threatens the scope of both families. 

The affinities of Pachycordyle, or its synonyms Clavopsella and Thieliana, are 
disputed even by contemporary authors. Some authors, including Millard (1975), 
Bouillon (1985), and Calder (1988a), place Pachycordyle in the Bougainvilliidae, 
while Moni (1980) and Stepanjants et al. (2000) firmly associate it with Cordylophora 
and the Clavidae sensu Bouillon (1985). To the present author its affinities with the 
Bougainvilliidae appear to be more evident. Pachycordyle navis and Pachycordyle 
pusilla sometimes approach the bougainvilliid species Aselomaris michaeli Berrill, 
1948 so much, that they could be easily confused. Also, the hydranths of the genera 
Parawrightia Warren, 1907, Velkovrhia Matjasic & Sket, 1971 or Pruvotella Motz- 
Kossowska, 1905 are intermediate between Pachycordyle and Bougainvillia. All of 
them have two or more whorls of tentacles. Even Bougainvillia muscus (Allman, 
1863), certainly a typical representative of the Bougainvilliidae, may have two whorls 
of tentacles, although quite closely approximated (own observations on living 
Mediterranean material). But most importantly, Silhouetta uvocarpa Millard & 
Bouillon, 1973 has a trophosome identical to that of Pachycordyle, but it produces 
medusae with branched oral tentacles. This definitely indicates a closer relationship to 
the Bougainvilliidae. The occurrence of nematocysts on the eggs of Pachycordyle 
navis (see Thiel, 1962, as Clavopsella quadranularid) and in many other bougainvil- 
liids as well (Russell, 1953 for B. macloviana; Szollosi, 1969 for B. multitentaculata; 
Calder, 1971 forß. rugosa; Piraino, 1992 for A. michaeli; Calder 1993 for B. abberans; 
Schuchert, 1996 for several species) could be interpreted as a synapomorphy. However, 
the distribution of such "stinging eggs" (Piraino, 1992) among the Filifera is too in- 
sufficiently known to draw a well supported conclusion. The few Oceanidae examined 
for this study apparently lack a conspicuous nematocyst layer on their egg surface. 



350 P. SCHUCHERT 

Contrary to the view of Calder (1988a), the genus Millardiana is not included 
in the Pachycordylidae because it develops gonophores on the hydranth body (see 
under family Oceanidae, discussion relating to Tubiclava). 

The genus Silhouetta is thus far not known to occur in the region under inves- 
tigation, but Cornelius (1992) recorded Silhouetta uvocarpa Millard & Bouillon, 1973 
from the nearby Azores. Considering its wide occurrence, ranging from the Seychelles 
to the western tropical Atlantic (Calder, 1988a), it might also occur along the coasts of 
southern Portugal or Spain. 

Pachycordyle Weismann, 1883 

Synonyms - Clavopsella Stechow, 1919; Thieliana Stepanjants et al, 2000. 
Type species - Pachycordyle napolitana Weismann, 1883 by monotypy. 

Diagnosis - Stolonal or branching hydroid colonies, hydranths pedicellate, 
pensare terminating at base of hydranth. Hydranths club-shaped, hypostome dome- to 
nipple-shaped, tentacles in two or more close whorls. Gonophores borne on stems or 
pedicels. Gonophores fixed sporosacs or degenerated pyriform medusae lacking 
mouth, marginal tentacles, oral tentacles, radial canals, or ocelli. 

Remarks - Clavopsella Stechow, 1919 is clearly a junior synonym of Pachy- 
cordyle Weismann, 1883 (Calder, 1988a; Stepanjants et al., 2000). The definition of 
Pachycordyle adopted in this work follows that used by Thiel (1962) and Millard 
(1975) for Clavopsella. This definition also includes Pachycordyle navis and other 
similar species. Calder (1988a) did not agree with this and considered P. navis as 
belonging to the subfamily Bougainvilliinae rather than the Pachycordylinae and he 
referred Pachycordyle navis to the genus Aselomaris Berrill, 1948. Stepanjants et al. 
(2000) again placed P. navis in the same subfamily as Pachycordyle napolitana, but in 
the new genus Thieliana. Thieliana differs from Pachycordyle only in the degree of 
reduction of its gonophores. While Pachycordyle sensu Stepanjants et al. (2000) has 
either radial canals or a circular canal, they are absent in Thieliana. In the remarks 
relating to Pachycordyle napolitana, I will show that there is probably no real circular 
canal present in this species. The radial canals of Pachycordyle kubotai Stepanjants et 
al., 2000 are additionally somewhat unusual in that they are not between layers of 
epidermis. Perhaps they are better homologized with a branched spadix. Generic 
distinction based on the degree of gonophore reduction have been shown to lead to 
artificial taxa in many cases (e. g. Petersen 1990, Bouillon et al. 1997 ', Boero et al. 
1998, Schuchert 2001b). The degree of gonophore reduction, notably if it is only so 
slight as in the case for Thieliana, should therefore not be used to distinguish genera if 
this trait does not concord with other, independent characters. Thieliana is here thus 
treated as a synonym of Pachycordyle. 

Kramp (1959, 1968) allocated a number of degenerate medusae to Pachy- 
cordyle. Kramp himself considered these medusae as possibly belonging to very 
different taxa and they are placed only temporarily in Pachycordyle pending forth- 
coming information on their polyp phase. 

In the region under investigation, there are three known species referable to the 
genus Pachycordyle: P. napolitana, P. navis, and P. pusilla. Piraino (1992) reported 



OCEANIDAE AND PACHYCORDYLIDAE 351 

Clavopsella michaeli (Berrill, 1948) from the Mediterranean. This species has a single 
whorl of tentacles and is here seen as belonging to the Bougainvilliidae. It should the- 
refore not be included in the genus Clavopsella, or its senior synonym Pachycordyle. 

Pachycordyle napolitana Weismann, 1883 Fig. 11 

Pachycordyle napolitana Weismann, 1883: 87; Neppi, 1921: 5, fig. 1; Calder, 1988a: 15, figs 

11-12. 
Cordylophora annulata Motz-Kossowska, 1905: 66, fig. 5; Stechow, 1923: 56. 
Perigonimus neapolitanus - Motz-Kossowska, 1905: 75, fig. 8 [incorrect subsequent spelling], 
[not Perigonimus napolitanus Hargitt, 1904 = Leuckartiara octona (Fleming, 1823)]. 
Pachycordyle weismanni Hargitt, 1904: 553, pi. 21 figs 1-8; Kramp, 1959: 94; Kramp, 1961: 52. 
Tubiclava annulata - Stechow, 1912: 343, pi. 13 fig. 8. 
? Tubiclava fruticosa - Neppi, 1917: 47, fig. 9. 
Clavopsella weismanni - Stechow, 1923: 55. 
Clavopsella annulata - Wedler & Larson, 1986: 86, fig. 8Aa-f, 9. 
? not Cordylophora neapolitana - Morri, 1980: 159, figs 4-5; Morii & Boero, 1986: 34, fig. 15c; 

Moni, 1981: 47, fig. 14, pi. 1 fig. 4 [incorrect subsequent spelling]. 

Material examined - IRSNB, Italy, Naples, young female colony collected by J. Bouillon, June 
1962 - ZSM, Stechow collection, as Clavopsella weismanni several colonies from 
Naples collected 1897-1904 as alcohol preparation and slide material, includes male and 
female colonies, gonophores of variable developmental stages, all colonies on Fusinus 
rostratus. 

Type material - There presumably exists no type material of P. napolitana. 

Description - Colony stolonal, or rarely pedicels branched once. Pedicels of 
very varied length (1-15 mm, usually below 5 mm), usually widening distally, bearing 
a terminal hydranth. Perisarc moderately thin, irregularly corrugated throughout, not 
distinctly double-layered, terminating at base of hydranth, not forming pseudohy- 
drotheca. Hydranths club-shaped, 1-2 mm long, diameter about 0.3 mm if expanded, 
when contracted much thicker, in distal third 12-24 tentacles, in larger hydranths 
distinctly scattered in a broad band (3 whorls), in smaller hydranths more concentrated 
into a narrow band and almost appearing as one whorl only, proximal tentacles often 
smaller than distal ones, contracted tentacles filiform, well expanded tentacles monili- 
form. Hypostome dome- to nipple-shaped. Gonophores arise from hydranth pedicels at 
some distance below hydranth, 1-3 per pedicel, on short stalk, completely infested with 
soft, wrinkled, detritus-covered perisarc, shape ovoid, length up to 1.6 mm, diameter 
0.8-1 mm. Mature gonophore medusoid, with a distinct subumbrella and a short collar 
projecting into subumbrella resembling a velum (Fig. 111). Radial canals absent, 
circular canal presumably absent. Vestigial manubrium simple, large. Below manu- 
brium a bubble-like gastrodermal chamber (Fig. 11G-I). Gonads encircling manubrium 
in a thick layer so that it is completely covered. Females with 100-150 eggs, oocytes 
with relatively large pronucleus, eggs shed and developing free in the water. Egg size 
about 0.07 mm. Immature male gonophores resembling sporosacs, manubrium large 
and with pointed diverticulae, mature male sporosacs also medusoid and identical to 
female ones. 

Further Data - Shoots may have up to three hydranths (Weismann, 1883). 
Hydranth body orange or reddish, hypostome whitish (Hargitt, 1904). Nematocysts are 
microbasic euryteles and desmonemes (Calder, 1988a). 



352 



P. SCHUCHERT 




Fig. 11. Pachycordyle napolitana Weismann, 1883, after preserved material from Naples. A) 
Hydranth with somewhat contracted tentacles, scale equals 0.4 mm. B) Hydranth, note that 
tentacles are less scattered than in specimen shown in A, same scale as A. C) Tip of a fully 
expanded tentacle, note moniliform arrangement of nematocyst clusters, scale equals 50 /<m. D- 
F) Successive developmental stages of a male gonophore, note branched spadix (manubrium), 
scale equals 0.3 mm. G) Fully mature male medusoid, it is still enveloped by perisarc film, same 
scale as D. H) Mature female medusoid, note that there is no more tissue connection to the polyp 
stage, but the medusoid is still within the perisarc envelope, same scale as D. G) Higher magni- 
fication of the distal opening of the female medusoid, ve = velum-like collar, cl = cleft, a possi- 
ble fixation artefact and likely not a circular canal, scale equals 0.1 mm. 



Biology - Reliable identifications are from fully marine conditions only. 
Colonies usually grow on gastropod shells (Fusinus rostratus, Murex spec), either 
inhabited by mollusc or hermit crabs, mostly on muddy bottoms; known to occur in 
depths from a few metres to perhaps 40 m; mature from spring to October. Boero & 
Fresi (1986) also found it growing on algae and Eudendrium spec, (fully marine 
conditions). 



OCEANIDAE AND PACHYCORDYLIDAE 353 

Distribution - Western Mediterranean (Weismann, 1883; Hargitt, 1904; Motz- 
Kossowska, 1905; Lo Bianco, 1909; Stechow, 1919; Neppi, 1921; Stechow, 1923; 
Boero & Fresi, 1986), Adriatic Sea (Stechow, 1912), Bermuda (Calder, 1988a), Puerto 
Rico (Wedler & Larson, 1986). Type locality: Mediterranean, Naples, 40 m, on empty 
Murex shell. 

Remarks - Weismann (1883) described P. napolitana only summarily, as he 
was more interested in its histology and in the development of the gonophores. The 
gonophores he described were male, appeared like sessile sporosacs, and had a 
characteristic, branched spadix. Hargitt (1904), also studying material from Naples, 
found similar colonies which were all female. Although he noted the close resemblance 
to P. napolitana, he assigned his material to a separate species and named it P. weis- 
manni, based mainly on the medusoid gonophores he observed. Shortly afterwards, 
Motz-Kossowska (1905) published her findings of athecate hydroids from the western 
Mediterranean. Motz-Kossowska, most probably unaware of Hargitt's publication, 
also described a new species with several whorls of filiform tentacles and female 
medusoid gonophores and named it Cordylophora cumulata. A similar, male colony 
with sporosacs identical to Weismann 's material was allocated by her to P. napolitanus 
under the name Perigonimus neapolitanus (Weism.) (not to be confounded with P. 
neapolitanus Hargitt, 1904, presumably a synonym of Leuckartiara octona). Mayer 
(1910) concluded that P. napolitana and P. weismanni belonged to the same species 
and that the differences of the two nominal species were attributable only to differences 
of the sex. Neppi (1921) put forward the same thoughts and substantiated this by 
observations. She demonstrated that the male gonophores of P. napolitana are also 
medusoid and have intermediate developmental stages resembling those described by 
Weismann. Stechow (1923) vehemently rejected the idea of Mayer and insisted that P. 
napolitana and P. weismanni are separate species. Additionally, he also kept C. annu- 
lata distict from P. weismanni, although without giving arguments. A re-examination 
of Stechow's material, today kept by the ZSM, makes it difficult to understand 
Stechow 's opinion, as the material clearly shows several developmental stages of male 
gonophores (Fig. 11D-G). It is especially important to note the development of the 
spadix, which initially has very characteristic lateral diverticulae, resembling a 
branched spadix (Fig. 11F). Such a spadix was also described by Weismann (1883). 
There can thus be no doubt that C. napolitana and P. weismanni are conspecific, 
although they were initially recorded on different gastropod shells {Murex spec, versus 
Fusinus rostratus). Thiel (1962) continued to use P. weismanni, and he kept P. annu- 
lata Motz-Kossowska, 1905 separate on account of the absence of a circular canal and 
the different branching degree, although Motz-Kossowska did not explicitly mention 
the absence of such a ring canal. Hargitt (1904) made histological sections of the 
medusoids and found a cleft-like space encircling the bell opening which he interpreted 
as a circular canal, although it lacked an epithelial lining. In the slide material made by 
Stechow, a similar cleft can be seen (Fig. Ill), but it is very likely that it is only a 
fixation artifact as in Hargitt's material. Thus, Pachycordyle napolitana very likely has 
no real circular canal. 

Pachycordyle napolitana has usually been reported growing on gastropod shells 
in waters of normal salinity, both in the Mediterranean and elsewhere (Wedler & 



354 P. SCHUCHERT 

Larson, 1986, as C. annulata; Calder, 1988a). Boero & Fresi (1986) recorded it also on 
algae and Eudendrium spp., also in a fully marine environment. Moiri (1980, 1981) 
reported P. napolitano, on seagrasses growing in brackish waters. It seems, however, 
that medusoid gonophores were never observed in these animals (see comment in 
Moni, 1980: 163). Because this is an essential character in distinguishing them from 
other similar species, like P. navis (Millard, 1959) (see Stepanjants et al., 2000), I think 
that the identity of Morn's material is not reliably established and needs confirmation. 
The difference of biotopes - fully marine versus reduced salinity and different 
substrata - are also arguments for two different species being involved. Pachycordyle 
navis is quite similar and is well known to occur in brackish waters. 

As already noted by Stechow (1923), the tentacles in fully grown hydranths of 
P. napolitana are clearly arranged in several whorls, but this may not be the case in 
smaller or younger hydranths. In preserved and contracted material it can also be quite 
difficult to recognize more than one tentacle whorl. 

Hargitt (1904) observed in aquarium cultures that the mature gonophores are 
released as free medusoids. As noted by Stechow (1923), this may not necessarily be 
the case under natural conditions, and some gonophores may spawn even while still 
attached to the hydranth. 

Pachycordyle navis (Millard, 1959) comb. n. Fig. 12 

Rhizorhagium navis Millard, 1959: 244, fig. 2. 

Clavopsella quadranularia Thiel 1962: 228, figs 1-28; Schönborn et al., 1993: 217, pi. 2 fig 3. 

Corydendrium dispar - Rasmussen, 1973: 22. 

[not Corydendrium dispar Kramp. 1935]. 

Clavopsella navis - Millard, 1975: 100, fig. 3A-D. 

? Cordylophora neapolitana - Moni, 1980: 159, figs 4-5; Morri & Boero, 1986: 34, fig. 15c; 

Morri, 1981: 47, fig. 14, pi. 1 fig. 4. 
[not Pachycordyle napolitana Weismann, 1883]. 

Cordylophora inkermanica Marfenin, 1983: 1732, fig'd; Stepanjants et al. 2000: 226. 
Thieliana navis - Faasse & Vervoort, 2001: 181, figs 1-3. 

Type material examined - Type of Clavopsella quadranularia Thiel, 1962, Zoological 
Museum Hamburg, registration number C7434, loc. Kiel, Aussenhafen, Holtenauer 
Schleuse. The type material of P. navis is kept by the South African Museum in 
Capetown, but was not examined during this study. 

Other Material examined - Material described by Rasmussen (1973) as Corydendrium dispar, 
Denmark. Isefjord. on Mytilus in the strong currents of the intake of the power station 
Kyndbyvaerket, fertile female, material kept by "The Isefjord Laboratory" at Vellerup 
Vig (University of Roskild. Denmark), part of this material also deposited as MHNG 
INVE 34226. 

Description - (After Thiel, 1962 and examined material) Colony stolonal or 
variably branched, height 7-30 mm, 1-30 hydranths per shoot. Stolons ramified, 
initially creeping on substrate, in larger colonies detached and forming tangled mass- 
es, sometimes also growing along stems of older shoots. Pedicels and stems covered 
by perisarc, terminating below hydranth. Perisarc mostly smooth, some annulated 
stretched in European colonies. Perisarc often double-layered, inner layer may be 
annulated while outer layer is smooth (Fig. 12C). Pedicel length variable but relatively 
long, diameter 0.1-0.15 mm. Hydranth body 1.0-1.8 mm high (1.4 mm mean), diameter 
about 0.35 mm. spindle-shaped. Hypostome high, dome-shaped. Tentacles 12-24, in a 



OCEANIDAE AND PACHYCORDYLIDAE 



355 




psi pso 



eoe 



Fig. 12 
Pachycordyle navis (Millard, 1959), A-B, after type material of Clavopsella quadranularia; C, 
after material from Denmark. A) Hydranth with part of pedicel, scale equals 0.2 mm. B) Female 
sporosac, scale equals 0.2 mm. C) Higher magnification of double-layered perisarc, eoe = 
coenosarc, psi = inner perisarc layer, pso = outer perisarc layer, note the thin lamellae connecting 
both layers, scale equals 50 /<m. 

narrow band below hypostome, 2-4 closely approximated alternating whorls. 
Gonophores about 0.4 mm, borne in irregular spiral on stem or hydranth pedicels. 
Gonophores with relatively long pedicel, completely sheathed in perisarc. Gonophores 
fixed sporosacs (heteromedusoids) with distinct spadix, lacking tentacle rudiments, 
radial canals, circular canal, or velum. Colonies dioecious. Male sporosacs spherical to 
spindle-shaped. Female sporosacs club-shaped, distal end obtuse, containing a very 
variable number of eggs, 5-20, if up to 12 eggs then these in one tier, but arrangement 
can also be irregular, eggs about 0.1 mm in diameter, covered with a few nematocysts 
only. Eggs develop into planulae in situ. Nematocysts: microbasic euryteles, 
desmonemes. Colour: perisarc clear to brown, coenosarc white to reddish, hydranths 
red (depends on food). 

Biology - Euryhaline. European colonies known from brackish waters only, 
the minimal salt concentration is perhaps 8 %c\ experimental cultures in seawater of 
normal salinity kept well over long time (Thiel, 1962). The South African colony was 
growing on a ship hull that had never left South Africa. Grows on algae, wood, iron 
constructs, Mytilus, and other solid substrata. Thiel (1962) observed two periods of 
reproductive activity: in spring and late summer to autumn. 

Distribution - South Africa (Millard, 1975), Baltic Sea (Thiel, 1962, as C. 
quadranularia), southern England (Barnes, 1994), The Netherlands (Faasse & 
Vervoort, 2001), Denmark (this study), Black Sea (Marfenin, 1983, as Cordylophora 
inkermanica). Type locality: Table Bay, South Africa, on hull of ship. 



356 P. SCHUCHERT 

Remarks - This species was initially described as Rhizorhagium navis Millard, 
1959. Later Millard (1975) placed it in the genus Clavopsella. Because Pachycordyle 
Weismann, 1883 has priority over Clavopsella Stechow, 1919, this species is here 
referred to the former genus as Pachycordyle navis (Millard, 1959) new comb. 

Millard (1975) reported that she had compared material off. navis with Thiel's 
Clavopsella quadranularia and both authors agreed that they were very likely conspe- 
cific. 

As put forward by Stepanjants et al. (2000), Cordylophora inkermanica 
Marfenin, 1983 appears indistinguishable from P. navis and it is here regarded as a syn- 
onym of the latter. 

Material identified by Rasmussen (1973) as Corydendrium dispar was re- 
examined during this study. Although not well preserved, it is obviously referrable to 
P. navis. Some preserved hydranths have the typical tentacle arrangement of this 
species. The side branches arise at acute angles, but remain not adnate as in C. dispar. 
Also, the well preserved female sporosacs were rather typical. The double-layered 
perisarc seen in both C. dispar and P. navis may have mislead Rasmussen to identify 
his material as C. dispar. 

Pachycordyle navis resembles P. pusilla Motz-Kossowska, 1905, but the latter 
is smaller, is stolonal, has fewer eggs per sporosac, and has a funnel-shaped perisarc 
dilation at the distal end of the pedicel. Furthermore, their ecology is different, with P. 
pusilla occuring only on sea-grasses in fully marine environments. European records 
of P. navis are all from sites with reduced salinities. There is an interesting similarity 
in the structure of their perisarc (comp. figs. 12C and 13D). The perisarc is often 
doubled with a thicker, corrugated inner layer and a thin outer layer. Both layers are 
connected by thin lamellae. In P. pusilla, this double layered-structure is confined to 
the distal part of the pedicels. 

Pachycordyle pusilla (Motz-Kossowska, 1905) comb. n. Figs 13 & 14 

Cordylophora pusilla Motz-Kossowska, 1905: 63, fig. 4, pi. 3 fig. 3-9; Moni, 1980: 164, fig. 6; 
Boero, 1981: 188, fig. 2; Schlichen 2003a: 540, fig'd. 

? Pachycordyle fusca Müller, 1913: 357, figs 3-7, figs 14-23. 

Tubiclava pusilla - Stechow, 1919: 10; Leloup, 1930: 24, figs 1-3. 

Merona pusilla - Picard, 1951: 349. 

Type material examined - ZSM, syntype material of Pachycordyle fusca, Stechow collection, 
one slide labelled as "Clavopsella {Pachycordyle) fiasca (H. G. Müller) Neapel Cotypus 
!!" (cotypus in red ink), 2 hydranths with young gonophores (stage 1 acc. Müller, 1913: 
399), gastrodermal zooxanthellae clearly visible. The type material of P. pusilla could 
not be located. 

Other Material examined - IRSNB, Mediterranean, Monaco, coll. E. Leloup 1929, on 
Posidonia oceanica (material of Leloup, 1930) - IRSNB, Mediterranean, Villefranche- 
sur-Mer, coll. E. Leloup 24 Mar 1934, depth 15 m, on Posidonia oceanica - ZSM, 
Naples, as Clavopsella fusca, coll. Bedot no. 186, on Cymodocea nodosa, with young 
gonophores - ZSM, as Clavopsella fusca. Naples, coll. Splettstósser, fertile females - 
MHNG INVE 32953, Banylus-sur-Mer. on Posidonia oceanica, 10 May 2002, examined 
alive, fertile female. 

Description - Colony stolonal or rarely branched once, stolons creeping, 
smooth, mostly linear, growing on seagrasses. Hydranths pedicellate, pedicel 0.4-2 mm 
high, with perisarcal sheath, perisarc annulated in lower region, at distal end often 



OCEANIDAE AND PACHYCORDYLIDAE 



357 



> 1 


H 

fi 

il 

j,' 


\ it 
\ \\ 


ff / 


% ' ! 'l 


l'I /? 


■^N 'À l'i 


i' // 


\ \\\ 


// // 


1 v v^, 








Fig. 13 
Pachycordyle pusilla (Motz-Kossowska, 1905). A) hydranth with female sporosac, drawn after 
living material from Banyuls-sur-Mer, scale equals 0.4 mm. B) Hydranth with contracted 
tentacles, note scatter of tentacles, preserved material from Monaco, scale equals 0.4 mm. C) 
Optical section of hydranth pedicel, note double-layered, lamellar structure of distal end, same 
scale and material as in B. D) Optical section of hydranth pedicel with young female sporosac, 
preserved material from Naples, scale equals 0.2 mm. E) Final developmental stage of female 
sporosacs with three fully developed planulae, same material as D, scale equals 0.2 mm. F) Male 
sporosac in optical section, preserved material from Villefranche, scale equals 0. 1 mm. 



358 



P. SCHUCHERT 





B 



Fig. 14 
Pachycordyle fiisca Müller, 1913, after type material. A) Hydranth, some tentacles are broken 
off, note their scattered arrangement, scale equals 0.2 mm. B) Higher magnification of pedicel 
perisarc in optical section, note annulation and double-layered periderm, the inner thicker and 
the outer layer are connected by lamellae, scale equals 50 /mi. 



expanding and fuzzy, double-layered, the two layers connected by lamellae (Figs 13D), 
sometimes resembling nested funnels. Hydranth 0.7-1.2 mm, fully relaxed hydranths 
slightly club-shaped with swelling in region of tentacles (Fig. 13A), hypostome high, 
dome-shaped. Tentacles radiating from a narrow band below hypostome, 12-16 in 
number, either in 3 whorls or scattered, expanded very thin, up to 1 .4 mm long. Size 
of tentacle-bearing region quite variable, in some hydranths very narrow, giving the 
impression of only one whorl being present. Gonophores borne on the hydranth 
pedicels, usually near the distal end, 1-4 per pedicel. Gonophores sessile sporosacs, 
without radial or circular canals, no velum, simple spadix present, proximal gastro- 
dermal chamber large, taking up almost the same volume as the gametes. Gonophores 
surrounded by loose, thin periderm. Females with 4, occasionally 5, eggs in a single 
tier, eggs uncoloured, clear. Development takes place in sporosacs, planulae remain 
attached to spadix even after periderm casing has been shed. Nematocysts: microbasic 
euryteles and desmonemes. 

Biology - Grows on Posidonìa oceanica or Cymodocea nodosa in sea water of 
normal salinity. Mature colonies found from June to August (Motz-Kossowska, 1905), 
March to April (Leloup. 1930), October (Boero & Fresi, 1986). Reproduction leads to 
reduction of hydranths (reproductive exhaustion; Leloup, 1930). 



OCEANIDAE AND PACHYCORDYLIDAE 359 

Distribution - Mediterranean. Records: Banyuls-sur-Mer (Motz-Kossowska, 
1905); Marseille (Stechow, 1919); Monaco (Leloup, 1930); Algeria (Picard, 1955); 
Genoa (Boero, 1981; Boero & Fresi, 1986); Cabo de Creus, Spain (Gili & Castello, 
1985). Type localities: Pachycordyle pusilla, near Arago Laboratory, Banyuls-sur-Mer, 
France, Mediterranean, on Posidonia oceanica; Pachycordyle fiisca, Bay of Naples, on 
Cymodocea nodosa, 2-4 m. 

Remarks - Contemporary authors placed this species in the genus 
Cordylophora. The tentacle arrangement, however, is identical to that of Pachycordyle 
napolitano and P. navis. All three species resemble each other quite closely, although 
they are certainly distinguishable. Therefore, this species is here used in the new 
combination Pachycordyle pusilla (Motz-Kosowska, 1905). 

Most hydranths have their tentacles scattered in a band below the hypostome, 
but in some hydranths they are so close that they can be taken as one whorl only. These 
polyps look very much like bougainvilliid hydranths. 

Motz-Kossowska (1905) mentioned that the eggs are covered by nematocysts. 
In the examined material from Banyuls, though, only very few nematocysts were 
present on the egg surface. 

Zooxanthellae could not be detected reliably in the examined living hydranths, 
but they may have been present in low numbers. 

Pachycordyle fusca Müller, 1913 very much resembles P. pusilla in all aspects, 
but its gastrodermal tissue contains many zooxanthellae. The zooxanthellae give the 
hydranth a brown colour. The species has apparently not been mentioned by other 
authors since its original description. Müller (1913) described P. fiisca quite precisely, 
especially the gonophores and their development. In one respect, however, he might 
have been wrong: the tentacles are not in a single whorl as shown in his figure (Müller 
1913: fig. 3). The ZSM collection contains a slide that is very likely syntype material 
because it is clearly labelled "Cotypus". It was presumably donated by Müller to 
Stechow, who initiated the hydroid collection of the ZSM (these authors must have 
been in contact because they also published together, e. g. Stechow & Müller, 1923). 
This slide preparation contains two hydranths that clearly match Müller 's figure, 
including the double layered perisarc (Fig. 14B). The tentacles, however, are clearly in 
three whorls (see Fig. 14A). The zooxanthellae are visible in the preparation. Besides 
the zooxanthellae, and perhaps the host plant, there is thus nothing that distinguishes 
P. fiisca reliably from P. pusilla, and I suspect that both are conspecific. Pachycordyle 
fusca is therefore listed as a questionable synonym of P. pusilla. 

Müller (1913) primarily investigated the development of the gonophores and 
the regeneration of the hydranths and gonophores. He also kept his colonies in the dark 
which considerably reduced the zooxanthellae, but which had no adverse effect on the 
hydroid. 

Picard (1955) identified hydroids as Cordylophora pusilla that were growing on 
Cymodocea. He noted that the gastrodermis contained a small number of zoo- 
xanthellae. 



360 P. SCHUCHERT 

ACKNOWLEDGEMENTS 

I owe many thanks to the Claraz foundation, which provided financial support 
for field work conducted for this study. Many thanks are also due to several museums 
(BMNH, IRSNB, RMNH, ZMUC, ZSM) for their generous loans of material. Dr S0ren 
A. Nielsen (University of Roskilde) kindly provided me with material of the 
Rasmussen collection. My collègues Dr F. Boero, Dr D. Calder, and Dr A. Marques 
took the burden to read and correct earlier version of this manuscript. Their construc- 
tive corrections and suggestions - although not all could be followed - significantly 
improved the quality of this publication. I wish to express my deep gratitude for their 
efforts. The production of the histological sections by Mrs Janik Pralong was also 
much appreciated. 

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Revue suisse de Zoologie 111 (2): 371-380; juin 2004 



Redescription of Ophiotaenia hylae Johnston, 1912 (Eucestoda: 
Proteocephalidea), parasite of Litoria aurea (Amphibia: Hylidae) 
from Australia 

Alain DE CHAMBRIER 

Muséum d'histoire naturelle, Département des Invertébrés, PO Box 6434, 
CH-1211 Geneva 6, Switzerland. E-mail: alain.dechambrier@mhn.ville-ge.ch 



Redescription of Ophiotaenia hylae Johnston, 1912 (Eucestoda: Proteo- 
cephalidea), parasite of Litoria aurea (Amphibia: Hylidae) from 
Australia. - Type material of the proteocephalidean cestode Ophiotaenia 
hylae Johnston, 1912 is redescribed. It is characterised by a globular scolex 
with uniloculate suckers, a prominent apical organ covered by spiniform 
microthriches and containing round to oblong cells of finely granular cyto- 
plasm, and by the internal longitudinal musculature composed by 4-5 dorsal 
and 4-5 ventral bundles of fibres. A similar taxon, Ophiotaenia sp. from a 
closely related host species, Litoria moorei, is also studied and compared. 

Key-words: Eucestoda - Proteocephalidea - Ophiotaenia hylae - Litoria 
aurea - Litoria moorei - Hylidae - Australia. 

INTRODUCTION 

Since its original description in 1912, nobody made a redescription of O. hylae. 
Prudhoe & Bray (1982, p. 33, figs 8a, b, c) only published drawings of Ophiotaenia 
hylae (BMNH 1968.4.19.1-15) from Litoria (Hyla) moorei, Cannington, Australia. 
This material has been examined and is here considered as belonging to Ophiotaenia 
sp. (see below) and not to O. hylae Johnston. New material of the true O. hylae was 
unavailable due to the extreme scarcity of its host, Litoria (Hyla) aurea. This species 
is actually considered threatened with extinction in Australia (Pyke et al., 2002; Pyke, 
2002). I had the opportunity to study Johnston's type material deposited in different 
Australian museums (Queensland Museum, Brisbane; South Australian Museum, 
Adelaide). This allowed me to redescribe this material, to add new information to the 
original description and clarify its taxonomic status. 

MATERIAL AND METHODS 

The worms, conserved in museum collection in alcohol, were stained with 
Weigert's haematoxylin solution, dehydrated in an ethanol series, cleared with Eugenol 
(clove oil), and mounted in Canada balsam. Pieces of strobila were embedded in 
paraffin wax, cross sectionned (thickness 12-15 pirn), stained with Weigert's haema- 



Manuscript accepted 15.10.2003 



372 A. DE CHAMBRIER 

toxylin and counterstained with 1% eosin B according to the method recently 
published by de Chambrier (2001). All measurements are given in micrometres unless 
otherwise stated. 

Abbreviations used in descriptions: x = mean, n = number of measurements, CV 
= coefficient of variability (%), OV = ovary width to proglottis width ratio, PG = 
position of the genital pore in relation to proglottis length (%), PC = cirrus pouch 
length to proglottis width ratio (%), JNT = Johnston original description; BMNH = 
Natural History Museum, London; MHNG = Natural History Museum, Geneva; INVE 
= Geneva Museum, Invertebrates Collection; SAM = South Australian Museum, 
Adelaide; QM = Queensland Museum, Brisbane. 

RESULTS 

Ophiotaenia hylae Johnston Figs 1-6 

Ophiotaenia hylae Johnston, 1912: 63. 
Batrachotaenia hylae\ Rudin, 1917: 366. 
Batrachotaenia hylae; Freze, 1965: 385. 

Type host: Litoria aurea (Lesson, 1829) (Amphibia: Hylidae). 

Material studied: Syntype material of Ophiotaenia hylae: 1 slide V 4141 (SAM 44141); 
S 689 (SAM 20689), four slides: a) 2 immature pieces, one with scolex; b) 1 immature piece, 
9 mm; c) 1 gravid proglottis, 1 mm, bad conservation state; d) 11 gravid proglottides, 15 mm. 
One immature specimen with a scolex, G 16/423, QM, from Hyla aurea, Sydney, NSW, 37°41 'S, 
144°40'E. Other material: In one separate box containing a lot of Johnston's original material, 1 
slide with 12 immature pieces, with one scolex, SAM 28407. 

Site of infestation: Intestine. 

Type-locality: Neighbourhood of Sydney, NSW, Australia. 

Redescription (based on syntypes and Johnston's original material) 

Proteocephalidea, Proteocephalidae. Testes, ovary, uterus diverticles in me- 
dulla, uterine stem cortical. In the whole mounted syntype material, one fragment 18 
mm long. Strobila acraspedote, anapolytic, consisting of 33 immature and mature 
proglottides (JNT = 60 mm long). Immature proglottides 540-635 long and 405-500 
wide, mature proglottides 695-750 long and 710-865 wide, gravid proglottides 1250- 
1540 long and 865-920 wide (Figs 3, 4). Tegument thick and wrinkled in mature 
proglottides. Presence of numerous small dorso-ventral muscles. 

Scolex 340-390 (JNT = 320) in diameter (Figs 1-2), covered by small dense 
microtriches about 1 long, suckers 130-135 (JNT =110) in diameter. Apical organ, 65- 
80 in diameter, covered by small dense spiniform microtriches 2-3 long (Fig. 2) above 
a network of small and poorly defined canals filled with a granular content, and ending 
beneath tegument surface. Presence of small retractor musculature at the margin of the 
apical organ (Fig. 2). Beneath the apical organ, a concentration of cells with a finely 
granular cytoplasm is present in two zones, one just beyond the apical organ and 
another made of twice bigger cells situated at the level of the suckers (Fig. 2). Longi- 
tudinal internal musculature dense, formed by 4-5 thick bundles of fibres on both 
dorsal and ventral sides (Fig. 5). 

Ventral and dorsal osmoregulatory canals between vitelline follicles and testes, 
crossing cirrus pouch at level of its two/third part (Fig. 3). Ventral canal, twice the 



REDESCRIPTION OF OPHIOTAENIA HYLAE 



373 







Figs 1-2 
Ophiotaenia hylae Johnston, 1912. Syntype, 20689 SAM. 1. Scolex, general view. 2. Detailed 
view of the apical organ region. Abbreviations: cl, small canals- filled with granular content; gc, 
gland cells; lm, internal longitudinal musculature; rm, retractor muscles; ro, rostellum-like apical 
organ; sm, spiniform microtriches (hooklets). Scale-bars: 1 = 200 /mi; 2 =100 firn. 



374 



A. DE CHAMBRIER 




Fig. 3 
Ophiotaenia hylae Johnston, 1912. Syntype, 44141 SAM, dorsal view of a mature proglottis. 
Scale-bar: 500 pim. 



diameter of the dorsal canal, with narrow secondary canals directed externally ven- 
trally. Testes 74-106 in number (x = 86, n = 12, CV =13%, JNT = numerous) in two 
dorsal field, with tendency to converge anteriorly and posteriorly, in one or two layers 
dorsally, not reaching laterally to vitelline follicles (Fig. 3), 35-60 in diameter. Testes 
15-22 preporal, 16-25 postporal and 38-53 aporal in number. Testes degenerated in 
gravid proglottides (Fig. 4). 

Genital pores irregularly alternating, opening between 44 and 55 % (n = 12, CV 
= 7%) of proglottis length. Small genital atrium present. Cirrus pouch pyriform, US- 
MS long (JNT 140). PC = 17-19% (x = 18%, n = 11, CV = 5%). Cirrus occupying up 
to 70% of cirrus pouch length. Evaginated cirrus covered by numerous minute spini- 
form microtriches, 2-3 long. Vagina anterior (38%) or posterior (62%) to cirrus pouch 
(JNT = antero-ventrally), with a small subterminal vaginal sphincter (Fig. 3). When 
anterior, passing ventrally to the cirrus pouch. Mehlis glands 60-85 in diameter. Vas 
deferens coiled, between base of cirrus pouch and median part of proglottis, rarely 
extending beyond body midline in mature and premature proglottides, extending 
anteriorly. 



REDESCRIPTION OF OPHIOTAENIA HYLAE 



375 




Figs 4-7 
4-6. Ophiotaenia hylae Johnston, 1912. Syntype material, 20689 SAM. 4. Scheme of dorsal view 
of a gravid proglottis showing the uterine diverticle development. 5. Scheme of the internal lon- 
gitudinal musculature at level of immature proglottides. 6. Scheme of a cross section showing 
the disposition of genital organs related to the internal longitudinal musculature. 7. Ophiotaenia 
sp. from Litoria moorei. BMNH 1968.4.19.1-15, eggs drawn in distilled water. Abbreviations: 
em, embryophore; lm, internal longitudinal musculature; oe, outer envelope; on, oncosphere; ov, 
ovary; te, testes; ut, uterus; vi, vitellaria; Scale-bars: 4 = 500 ptm; 5 = 250 /<m; 6 = no scale; 7 = 
20 pirn. 



Ovary bilobate, medullary, folliculate, with numerous dorsal outgrowths 
(Figs 3, 7). OV = 68-71% (x = 70%, n = 11, CV = 2%). Vitelline follicles, in two lateral 
bands, occupying porally 91-97% of proglottis length, and aporally 94-97% of pro- 
glottis length (Fig. 3). 



376 A. DE CHAMBRIER 

Primordium of uterine stem cortical, already present in immature proglottides, 
with diverticles in medulla. Formation of uterus of type 2 (see de Chambrier et al., 
2004): in immature proglottides, chromophil cells concentrated laterally on both sides 
of uterine stem; in the first mature proglottides, lateral ramified digitations without a 
lumen, occupying at this stage already about 35% of proglottis width; in gravid 
proglottides, lateral diverticles occupying up to 91% of gravid proglottis width. Uterus 
with 10-17 (JNT = numerous) lateral medullar ramified diverticles on each side (Fig. 
4) and one or sometimes several ventral apertures as described for Crepidobothrium 
spp (de Chambrier, 1989a, b). Eggs, measured in whole preparations, with oncosphere 
11-12 in diameter (JNT = 7.5-11), hooklets 5-6 long; embryophore 13-14 in diameter 
(JNT = 15-19); outer envelope 60-75 in diameter. 

Ophiotaenia sp. Figs 7-10 

Proteocephalus hylae; Prudhoe & Bray, 1982: 33, Figs 8a, b, c. 
[Not Ophiotaenia hylae Johnston]. 

Host: Litoria moorei (Copland, 1957) (Amphibia : Hylidae). 

Locality: Neighbourhood of Perth (Cannington and Darlington), W.A., Australia. 

Material studied: 9 whole mount preparations and material in alcohol (from where SEM 
microphotographs come from) ex Litoria moorei, Cannington, Western Australia, 17.04.1966: 
BMNH 1968.4.19.1-15; 1 whole mount preparation SAM 21402, Darlington, W.A., 12.11.1980. 

Site of infestation: Intestine. 

Description 

Proteocephalidea, Proteocephalidae. Testes, ovary, uterus diverticles in 
medulla, uterine stem cortical. Strobila acraspedote. anapolytic. Tegument thick and 
wrinkled in mature proglottides. Presence of numerous small dorso-ventral muscles. 

Scolex 260-340 (n = 4, x = 290) in diameter, suckers 105-130 in diameter 
(Fig. 9). Apical organ, 50-80 in diameter, covered by small dense spiniform micro- 
triches (Fig. 10). Longitudinal internal musculature dense, formed by 4-5 thick bundles 
of fibres on both dorsal and ventral sides (Fig. 8). 

Ventral and dorsal osmoregulatory canals crossing cirrus pouch at its middle 
part, situated between vitelline follicles and testes (Fig. 8). Ventral canal, twice the 
diameter of the dorsal canal, with numerous narrow secondary canals directed 
externally. 

Testes 46-76 in number (x = 59, n = 30, CV = 14%) in two dorsal field, with 
tendency to converge anteriorly and posteriorly, in one or two layers dorsally, not 
reaching laterally to vitelline follicles (Fig. 8), 50-80 in diameter. Testes 14-26 pre- 
poral, 6-18 postporal and 23-44 aporal in number. Testes degenerated in gravid pro- 
glottides. 

Genital pores irregularly alternating, opening between 46 and 57 % (n = 17, CV 
= 6%) of proglottis length. Small genital atrium present. Cirrus pouch pyriform, 175- 
215 long, PC = 27-33% (x = 29%, n = 17, CV = 4%). Cirrus occupying up to 85% of 
cirrus pouch length. Vagina anterior (54%) or posterior (46%) to cirrus pouch, with a 
sub-terminal vaginal sphincter (Fig. 8). When anterior, passing ventrally to the cirrus 
pouch. Mehlis glands 45-65 in diameter. Vas deferens coiled, between base of cirrus 
pouch and median part of proglottis, often extending beyond body midline in mature 
and premature proglottides, extending anteriorly. 



REDESCRIPTION OF OPHIOTAENIA HYLAE 



311 




Fig. 8 
Ophiotaenia sp. from Litoria moorei. 21402 SAM. Mature proglottis, dorsal view. Note the 
secondary canals ending beneath the tegument. Scale-bar: 500 }im. 



Ovary bilobate, medullary, folliculate, with dorsal outgrowths. OV = 55-63% 
(x = 60%, n = 17, CV = 3%) (Fig. 8). Vitelline follicles, in two lateral bands, occupying 
porally 87-91% of proglottis length and aporally 86-88% of proglottis length. 

Primordium of uterine stem cortical, already present in immature proglottides, 
with diverticles in medulla. Formation of uterus of type 2 (see de Chambrier et al., 
2004). 



378 



A. DE CHAMBRIER 









.: ^ ._:_.„ 



Figs 9-10 
Ophiotaenia sp., BMNH 1968.4.19.1-15, from Litoria moorei. Scanning electron micrographs 
of the scolex. 9. Dorsoventral view. 10. Apical view, detail of the apical organ. Scale-bars: 9 = 
50 firn; 10 = 10 pirn. 

Eggs, measured in distilled water, with oncosphere 12-18 in diameter, hooklets 
5-9 long; embryophore 18-23 in diameter; outer envelope up to 55 in diameter (Fig. 7). 

Remarks 

This taxon is similar to Ophiotaenia hylae on the basis of the following charac- 
ters: similar apical organ, position of the genital pore, presence of 4-5 dorsal and 4-5 
ventral longitudinal internal bundles of musculature. It differs from it by the number of 
testes (46-76 versus 74-106), by the cirrus pouch length/width of proglottis ratio (27- 
33% versus 17-19%), and by the ovary width / proglottis width ratio (55-63% versus 
68-71%). 

Although these observations suggest that it could belong to a new distinct 
species, the material studied is fragmented and not in suitable conditions for an 
accurate description. The scolex particularly is badly fixed. In order to confirm that it 
represents a new species, it would be necessary to collect new material from Litoria 
moorei. 



DISCUSSION 

Johnston (1912) situated the ovary and the vitelline follicles of Ophiotaenia 
hylae in the cortex. My observations show the ovary to be clearly medullary (see 
scheme, Fig. 6). As for the vitelline follicles, their position is difficult to assess as there 
are no clear lateral muscle bundles (Fig. 6). The uterus stem is cortical with further 
development of diverticles into the medulla (Fig. 6). Contrary to the opinion of 
Johnston (1912, p. 64), the uterus does not arise as "a thin duct..." but is clearly of the 
type 2 of uterine development as described by de Chambrier et al. (2004). According 
to Johnston (1912), the vagina is situated anterior and ventral to the cirrus pouch. I 
observed a position mainly posterior (62%) of the vagina. I also observed a small sub- 



REDESCRIPTION OF OPHIOTAENIA HYLAE 379 

terminal vaginal sphincter, secondary canals emerging from the ventral osmo- 
regulatory canal ending under the tegument and the position of vas deferens which 
extends anteriorly (see Fig. 3). 

The structure of the internal longitudinal musculature is also uncommon within 
the Proteocephalidea. It is composed by 4-5 dorsal and 4-5 ventral powerful isolate 
bundles of musculature easy to observe in immature and mature proglottides but less 
so in gravid proglottides. Given the stability of this character, the number of bundle in 
mature proglottides could be discriminant at the specific level as I already proposed for 
Crepidobothrium species (de Chambrier, 1989b, p. 369). 

The apical organ is peculiar because of the presence of small spiniform hooklets 
covering its surface, retractor-like muscles and network of small canals surrounding it. 
This morphology shows some similarities with that of the Gangesiinae and looks 
intermediate between the apical organs found in the Nomimoscolex piraeeba aggregate 
(Zehnder et al., 2000) and those in the Gangesiinae (de Chambrier et al., 2003). To my 
knowledge, no other Proteocephalidea have this kind of apical organ. It would be 
interesting to analyse the two Australian Ophiotaenia species described in the present 
paper using DNA sequences, and compare them with the taxa cited above in order to 
see if their respective apical organs could represent a possible evolutionary trend or if 
this structure is homoplastic. 

ACKNOWLEDGEMENTS 

I am indebted to Lester Cannon (Brisbane), Ian Whittington and Leslie Chilsom 
(Adelaide) for loan of the type material, David I. Gibson and Eileen Harris (London) 
for loan of the comparative material. I am deeply indebted to the "Donation Georges 
et Antoine Claraz" for supporting this study. I am grateful to Jean Mariaux (Genève) 
who provided fruitful comments on earlier versions of the manuscript, Jean Wuest 
(Geneva) for the SEM photography and to Gilles Roth (Genève) for his help with 
drawings. 

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viperis (Beddard, 1913). Revue suisse de Zoologie 96: 191-217. 

de Chambrier, A. 1989b. Révision du genre Crepidobothrium Monticelli, 1900 (Cestoda: Pro- 
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380 A. DE CHAMBR1ER 



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Program of Scientific Translation, 1969, Cat. No. 1853, v + 597 pp). 
Johnston, H. 1912. Notes on some Entozoa. Proceedings of the Royal Society of Queensland 

24: 63-91. 
Prudhoe, S. & Bray, R. A. 1982. Platyhelminth parasites of the Amphibia. British Museum 

(Natural History), London, 217 pp. 
Pyke, G. H. 2002. A review of the biology of the southern Bell frog Litoria raniformis (Anura: 

Hylidae). Australian Zoologist 32: 32. 
Pyke G. H., White, A. W., Bishop, P. J. & Waldman, B. 2002. Habitat-use by the green and 

golden Bell frog Litoria aurea in Australia and New-Zealand. Australian Zoologist 

32: 12. 
Rudin, E. 1917. Die Ichthyotaenien der Reptilien. Revue suisse de Zoologie 25: 179-381. 
Zehnder, M., de Chambrier, A., Vaucher, C. & Mariaux, J. 2000. Nomimoscolex suspectus n. 

sp. (Eucestoda, Proteocephalidea) with morphological and molecular phylogenetic 

analyses of the genus. Systematic Parasitology 47: 157-172. 



Revue suisse de Zoologie 111 (2): 381-393; juin 2004 



Chiave di determinazione dei Chirotteri (Mammalia) della 
Svizzera attraverso l'osservazione al microscopio ottico della 
struttura dei peli 

Riccardo PIERALLINI 1 , Albert KELLER 2 & Marco MORETTI 3 

1 Via Nosetto 4, CH-6987 Caslano, Svizzera. E-mail: pierallini.ric@bluewin.ch 

2 Muséum d'histoire naturelle, ite de Malagnou 1, CR 6434, CH-1211 Genève 6, 
Suisse. E-mail: albert.keller.plo6@bluewin.ch 

3 WSL Istituto federale di ricerca, Sottostazione Sud delle Alpi, CH-6504 Bellinzona. 

Identification key of bats (Chiroptera) from Switzerland based on hair 
structure observed through optical microscope. - The identification of 
mammalian hairs by microscope remains an efficient and well-known 
technique, but only a few studies concern bats. We studied hair morphology 
of the 29 species listed from Switzerland and present a dichotomic identi- 
fication key based on overhair morphology observed by using optical 
microscopes. This key allows to identify most of the genera and also some 
typical species. The final purpose of this key is to give the possibility to 
analyse and to identify quickly the hairs found in guano, taken from their 
bat roosts. 

Key-words: Chiroptera - hairs - optical microscope - identification key - 
guano - Switzerland. 

INTRODUZIONE 

L'osservazione al microscopio della morfologia dei peli permette di identificare 
gran parte dei mammiferi europei (Keller, 1978, 1980; Debrot et al, 1982; Teerink, 
1991). In questo contesto, tuttavia, gli studi sui Chirotteri non sono molto progrediti, 
sia per la mancanza di un interesse pratico, sia perché le conoscenze sul gruppo sono 
recenti (Benedict, 1957; Debrot et al, 1982; Keller, 1986; Teerink, 1991; Meyer et al, 
1997). I Chirotteri presentano una struttura dei peli particolare, che li distingue dagli 
altri mammiferi per la mancanza della parte midollare interna 3 , per contro le scaglie 
esterne della cuticola sono ben sviluppate e variamente strutturate, in special modo nei 
Microchirotteri (Benedict, 1957; Tupinier, 1973; Keller, 1986). Proprio osservando 
queste strutture singolari al microscopio ottico o elettronico, diversi autori già in 
passato avevano ventilato l'idea di classificare i Chirotteri sulla base della morfologia 
dei peli (Benedict, 1957; Tupinier, 1973; Dulie, 1978; Keller, 1986; Charvet & Keller, 



3 Fanno eccezione tre famiglie (Pteropodidae, Rhinopomatidae, Megadermatidae) nelle quali la 
parte midollare dei peli è presente (Benedict 1957). Essa è invece sempre assente nei Micro- 
chirotteri europei. 

Manoscritto accettato il 22.12.2003 



382 R- PIERALLINI ET AL. 

1989; Meyer et al, 1995), tanto che fu possibile arrivare a identificare diverse specie 
(Keller, 1986; Tupinier, 1973; Keller & Moeschler, 1988; Dove & Peurach, 2001). Fino 
ad oggi non era però ancora stata realizzata alcuna chiave di determinazione completa 
utilizzabile al microscopio ottico. 

Nel 1986, Keller elaborò una chiave di determinazione basata sulla struttura dei 
peli per le specie del genere Pipistrellus utilizzabile con l'ausilio del microscopio 
ottico. Tale strumento ha infatti il pregio di essere di facile uso e i preparati necessari 
non sono complessi da realizzare come quelli per l'analisi al microscopio elettronico 
(Charvet & Keller, 1989). Il presente studio vuole essere un seguito e un completa- 
mento di quel lavoro, presentando una chiave di determinazione che consideri le specie 
di Chirotteri presenti in Svizzera e consenta di identificare i vari generi (quando possi- 
bile anche le specie) a partire dall'osservazione al microscopio ottico della morfologia 
dei peli. 

Applicazioni pratiche della chiave di determinazione 

L'utilità pratica della chiave di determinazione presentata consiste nella possi- 
bilità che essa offre di analizzare e determinare i peli contenuti all'interno dello sterco 
dei pipistrelli (Charvet & Keller, 1989). I Chirotteri dedicano infatti parecchio tempo 
alla pulizia corporea e così facendo ingeriscono rilevanti quantità dei loro peli, che 
sono molto resistenti e non subiscono alterazioni durante il processo digestivo (Charvet 
& Keller, 1989). Diviene quindi possibile determinare gli animali a partire unicamente 
dallo sterco. 

In Svizzera la tutela dei pipistrelli è coordinata da due centri nazionali (CCO, 
KOF) 4 , cui fanno capo esperti e centri regionali che, per definire l'identità delle specie 
presenti nei rifugi (edifici, solai, grotte ecc.), effettuano spesso catture di animali. Nel 
Cantone Ticino il CPT 5 sta attuando un piano d'inventario che attribuisce ai rifugi una 
differente importanza in base alle specie presenti. L'esperienza ha mostrato che le 
catture provocano un notevole disturbo alle colonie e si rivelano solitamente laboriose. 
La possibilità di determinare il pelo contenuto nello sterco prelevato dai rifugi permette 
rilevamenti veloci e di vaste proporzioni, consentendo di orientare le azioni di cattura 
verso i rifugi o le aree di maggior interesse. Con questo metodo è inoltre possibile 
avere importanti indicazioni anche in assenza di animali, per esempio durante l'inverno 
o in rifugi abbandonati (presenza passata di specie oggi rare o estinte). 

Un altro aspetto utile della chiave di determinazione proposta consiste nella 
possibilità di esaminare il pelo di individui giovani: è infatti noto che la maggior parte 
dei criteri utilizzati per l'identificazione degli adulti (p. es. lo sviluppo del cranio, delle 
membra e dei denti) non è valida per i giovani. La struttura del pelo è invece la stessa 
sia nei giovani sia negli adulti (Keller & Moeschler, 1988). Neppure tra i sessi sono 
riscontrabili differenze (Benedict, 1957). 



4 CCO: Centre de coordination ouest pour l'étude et la protection des Chauves-souris, c/o 
Muséum d'histoire naturelle, CP 6434, CH-1211 Genève 6; KOF: Stiftung zum Schutze 
unserer Fledermäuse in der Schweiz, c/o Zoo Zürich, CH-8044 Zürich. 

5 Centro protezione Chirotteri Ticino, CH-6714 Semione. 



CHIAVE DI DETERMINAZIONE DEI CHIROTTERI 383 

MATERIALE E METODI 

La chiave di determinazione è stata realizzata mediante l'osservazione al micro- 
scopio ottico di peli prelevati da animali conservati in alcol o a secco nelle collezioni 
dei Musei di storia naturale di Ginevra (MHNG), Lugano (MCSN), Coirà (BNM), 
Barcellona (Spagna) e del Museo zoologico dell'università di Zurigo (coll. H. P. Stutz), 
nonché da animali catturati nel loro ambiente naturale durante altre indagini in corso 
nel Cantone Ticino (nessun animale è stato disturbato espressamente per la chiave). 
Sono state esaminate 29 specie caratteristiche della chirotterofauna Svizzera: 27 di 
queste sono citate in Hausser (1995) più due specie, Plecotus alpinus (Kiefer & Veith, 
2001) e Pipistrellus pygmaeus (Jones & Barratt, 1999), scoperte solo di recente, per un 
totale complessivo di 115 campioni (vedi "Materiale utilizzato"). 

La struttura dei peli delle diverse parti del corpo di un pipistrello è la stessa 
(Meyer et al., 1995). I peli analizzati sono stati prelevati principalmente dal dorso e 
dall'addome, di norma da individui adulti, anche se, come accennato, i giovani pre- 
sentano la stessa struttura del pelo (Keller & Moeschler, 1988). 

La preparazione dei campioni è stata eseguita col metodo descritto da Keller 
(1978, 1980, 1986) e Charvet & Keller (1989): i peli vengono puliti in xylol e succes- 
sivamente fissati su vetrino mediante balsamo del Canada. 

In generale su ogni pipistrello sono presenti in ordine sparso diversi tipi di peli: 
i peli lanosi, dalla struttura più semplice, e i peli primari e secondari, assai più 
strutturati. Per l'elaborazione della chiave sono stati utilizzati unicamente i peli secon- 
dari (come suggerito da Tupinier, 1973 e Keller, 1986). Essi sono composti dalla radice 
(che comprende il bulbo pilifero), dal fusto, che costituisce la gran parte della lun- 
ghezza del pelo (nella chiave esso è suddiviso in base, parte centrale e parte apicale) e 
infine dalla spatola, tipica appendice a forma di lancia (Fig. 1). 

Lo studio al microscopio ottico delle forme, delle dimensioni e delle peculiarità 
di ciascuna di queste parti del pelo ci ha permesso di confrontare e identificare i vari 
generi (e talune specie). Particolarmente importante è stata l'osservazione della dispo- 
sizione, della forma e dell'inclinazione che assumono le scaglie della cuticola (Fig. 2). 
In Benedict (1957), Tupinier (1973), Charvet & Keller (1989) sono descritti i diversi 
tipi di forme che esse possono assumere. Ai loro testi facciamo riferimento, quando 
parliamo di scaglie coronali o imbricate, che possono essere appiattite contro il fusto, 
divergenti o divaricate. Gli ingrandimenti usati per le osservazioni sono di 200, 400, e 
1000 volte. 

Al fine di agevolare l'utilizzo pratico della chiave, nelle note a pie di pagina 
sono integrate alcune informazioni sulle dimensioni dello sterco di talune specie. 
Infatti vi sono specie che, pur presentando una struttura dei peli assai simile, possono 
essere agevolmente differenziate sulla base di tali criteri (p. es. Myotis). 

Materiale utilizzato 

Barbastello, barbastellus (Schreber, 1774): BNM 13812, Andeer, GR; BNM 1602, S, 
Haldenstein, GR; MHNG 1709.071, Saillon, VS. 

Eptesicus nilssoni (Keyserling & Blasius, 1839): BNM 11673, Tinizong, GR; BNM 
9866, S, Brusio, GR; MHNG 1804.098 Fully, VS. 



384 



R. PIERALLINI ET AL. 



S 



1A IB 



IC 



Figs 1-2 
1: Morfologia generale dei peli: A, peli primari; B, peli secondari; C, peli lanosi, r, radice; f, 
fusto; s, spatola. 2: Morfologia delle scaglie della cuticola dei peli secondari: A, viste di profilo; 
B, viste di fronte. 



Eptesicus serotinus (Schreber, 1774): MHNG 722.35, F, Duillier, VD; MCSN VT 3202, 
M, Lugano, TI; MCSN VT 2614, FS, Cugnasco; MCSN VT 2415, M, Ascona, TI; MCSN VT 
2631, M, Cevio. 

Hypsugo savii (Bonaparte, 1837): MCSN VT 2662, F, Balerna, TI; MCSN VT 3211, F, 
Biasca, TI; MCSN VT 2661, F, Taverne, TI; MCSN VT 3209, M, Aquila, TI; MB 82.6812, 
Barcelona, Spagna. 

Miniopterus schreibersì (Natterer in Kuhl, 1819): MHNG 1492.81, M, Kephalovrysion, 
Argolis, Grecia; MHNG 949.54, M, Otrante, Italia; MCSN VT 17, Lugano, S.Martino, TI. 

Myotis bechsteini (Natterer in Kuhl, 1818): MZUZ 5911, Aedermanndorf; MZUZ 4717, 
Rechthalten; MZUZ 194, F. Wölflinswil; 

Myotis blythi (Tomes, 1857): MCSN VT 2583, F, Gordevio, TI. 

Myotis brandti (Eversmann, 1845): MHNG 1805.049, F, Geschinen, VS; MHNG 
1684.071, M, Boudry, NE. 

Myotis capaccinii (Bonaparte, 1837): MHNG 679.089, Lugano, TI; MHNG 728.052, TI; 
MHNG 968.001, Massât, Ariège, France. 

Myotis daubentoni (Leisler in Kuhl, 1819): MHNG 1684.064, Collonge-Bellerive, GÈ; 
MCSN VT 2573, M, Balerna. TI; MCSN VT 2629. M, Locamo, TI; MCSN VT 3217, F, Quinto, 
TI; MCSN VT 2665, FS, Caslano, TI; MCSN VT 16, Melide, TI. Ssp. nathalinae: MHNG 
1326.033 (paratype). F, Cabezarrubias, Spagna. 



CHIAVE DI DETERMINAZIONE DEI CHIROTTERI 385 



Myotis emarginatus (Geoffroy, 1806): MHNG 1071.014, M, Deluze, Doubs, France; C, 
F, Brusino, TI; MHNG 722.030, F, Valavran, GÈ. 

Myotis myotis (Borkhausen, 1797): MHNG, M, Crans/Céligny, GÈ; MCSN V 9, Pavia, 
Italia; MCSN VII; MCSN V 10, Pavia, Italia; MCSN V 8, Pavia, Italia; 

Myotis mystacinus: MHNG 1684.65, M, Roggwil, BE; C, M, Campo Vallemaggia, TI; 
MHNG 968.79, M, Champéry, VS. 

Myotis nattereri (Kuhl, 1818): MHNG 1714.045, M, La Ferrière, BE; MZUZ 1318, 
Feldis-Veulden ; MHNG 1120.031, M, Vevey, VD. 

Nyctalus lasiopterus (Schreber, 1780): MHNG 1010.056, Champéry, VS. 

Nyctalus leisleri (Kuhl, 1818): MCSN RC 43; MCSN VT 3210, F, Cornano, TI; MCSN 
VT 3231, F, Someo, TI; MCSN VT 2642, F, Minusio, TI; MCSN VT 2587, Losone, TI. 

Nyctalus noctula (Schreber, 1774): MHNG 1702.07, Genthod-Bellevue, GÈ; BNM 
5915, Felsberg. 

Pipistrellus kuhli (Natterer in Kuhl, 1819): PZ 434, M, Verona, Italia; MCSN VT 2612, 
Roveredo, GR; MCSN VT 2616, Chiasso, TI. 

Pipistrellus nathusii (Keyserling & Blasius, 1839): MB 82.6808, St.Pere Vilamajor, 
Spagna; MCSN VT 2478; MCSN VT 2648, Ascona, TI. 

Pipistrellus pipistrellus (Schreber, 1774): C, MS, Prugiasco, TI; C, Pura, TI; C, Monte, 
TI; C, Villaluganese, TI; C, Lavorgo, TI. 

Pipistrellus pygmaeus (Leach, 1825): MB 82.6809, M, Barcelona, Spagna; MHNG 
1828.017, M, Genève; C, Locamo, Bosco Isolino, TI; MHNG 1807.089, M, Mte Trodos, Cipro; 
C, Mte Generoso, TI; MHNG 1826.028, Genève; C, TI; C, Locamo, Bosco Isolino, TI; C, TI; C, 
Locamo, Bosco Isolino, TI. 

Plecotus auritus L., 1758: MHNG 1115.12, M, Ibach, SZ; MCSN VT 3214, F, Nante, TI; 
MCSN VT 20, Induno, TI; MCSN VT 3206, M, TI; C, M, Ambii, TI. 

Plecotus alpinus Kiefer & Veith, 2001: MHNG 1325.81, M, Lienz (Austria). 

Plecotus sp. (austriacus (Fischer, 1829)/ 'alpinus): C, M, Campo Vallemaggia, TI; C, M, 
Crana, TI; C, M, Salorino, TI. (animali catturati nel 2001 e subito rilasciati, la cattura è avvenu- 
ta prima della scoperta della nuova specie P. alpinus, Kiefer & Veith, 2001). 

Rhinolophus euryale (Blasius, 1853): MHNG 925.75, M, Les Baux de Provence, France; 
MHNG 905.091, M, Fort l'Ecluse, Ain, France; MHNG 905.092, F, Fort l'Ecluse, Ain, France. 

Rhinolophus ferrumequinum (Schreber, 1774): MHNG 1255.42, Tourtenay, France; C, 
Salorino, TI; MCSN VT 15, Carabbia, TI; MHNG 722.063, M, Andermatt, UR; MHNG 
890.009, M, Grotte du Poteux, VS; MHNG 976.073, M, Grotta del Tesoro, TI; MHNG 1255.045, 
M, Tourtenay, Deux-Sèvres, France. 

Rhinolophus hipposideros (Bechstein, 1800): MHNG 1255.49, M, Angliers, France; 
MCSN VT 39; MHNG 722.054, M, Baar, ZG; MHNG 722.055, M, Baar, ZG; MHNG 722.059, 
F, Satigny, GÈ; MHNG 903.033, M, Versoix, GÈ; MHNG 976.060, M, Genève; MHNG 
987.013, M, Pissevache, Vemayaz, VS; 

Tadarida tenions Rafinesque, 1814: MHNG 1065.090, M, Champéry, VS; MHNG 
946.001, M, Champéry, VS; MHNG 1044.013, M, Champéry, VS. 

Vespertilio murinus Linné, 1758: MHNG 852.30, F, GÈ; MHNG 1120.037, M, Vevey, 
VD; MHNG 949.13, M, Col de Bretolet, Champéry, VS; MHNG 1755.095, M, Genève. 

(MHNG = Museo di storia naturale di Ginevra; MCSN = Museo cantonale di storia natu- 
rale di Lugano; MZUZ = Museo zoologico dell'Università di Zurigo, coll. H.P Stutz; BNM = 
Museo di storia naturale di Coirà; MB = Museo di zoologia di Barcellona (Spagna); PZ = coli. 
P. Zingg; C = cattura di animali, dati presso CPT; M = maschio; F = femmina; S = subadulto). 

RISULTATI 

Chiave di determinazione delle famiglie 

1 II fusto forma una linea spezzata (Fig. 3A). La radice del pelo può pre- 
sentare un rigonfiamento (Fig. 4A). Il pelo è poco pigmentato . . Rhinolophidae 
Il fusto non forma una linea spezzata (Fig. 3B, C) 2 



386 



R. PIERALLINI ET AL. 




3A 




If iL-* '* 





.-- 



" - : *'* '**"*-/■' 



3D 



Fig. 3. Fusto. A: Rhinolophus hipposideros. B: Myoris myotis. C: Tadarida teniotis. D: Tadarida 
teniotis, base del fusto. 

2 Le scaglie del fusto sono asimmetriche e imbricate (Fig. 3B, 7 ). Pos- 

sono avere anche un aspetto coronale, tuttavia sono imbricate (Fig. 6A, 

B, C) Vespertilionidae 

Le scaglie del fusto hanno una struttura simmetrica, coronale e divari- 
cata (Fig. 3C). L'aspetto dei peli è molto caratteristico e uniforme. Alla 
base del fusto le scaglie formano strutture "a spine" (Fig. 3D) 
Molossidae: Tadarida teniotis 

Chiave di determinazione di generi e specie 
Rhinolophidae 

1 Presenza di un rigonfiamento nella radice del pelo (Fig. 4A) 2 

Assenza di un rigonfiamento nella radice del pelo (Fig. 4B) 
Rhinolophus euryale 

2 Sul fusto alcune scaglie sono coronali e formano ispessimenti (Fig. 4C) 

Rhinolophus ferrumequinum 

Sul fusto le scaglie non formano ispessimenti Rhinolophus hipposideros 

Vespertilionidae 

1 Sulla spatola le scaglie sono divergenti (Fig. 5 A, B, C) 2 

Sulla spatola le scaglie sono appiattite contro il fusto (fig. 5D, E) 7 

2 Sulla spatola le scaglie sono molto divergenti e fitte (Fig. 5A) 3 

Sulla spatola le scaglie sono meno divergenti e fitte (Fig. 5B, C) 6 



CHIAVE DI DETERMINAZIONE DEI CHIROTTERI 



387 




FlG 4. A: Rhinolophus hipposideros, radice. B: Rhinolophus euryale, radice. C: Rhinolophus 
ferrumequinum, ispessimento del fusto (l'ingrandimento è maggiore rispetto A e B). 




5A 



ìjK **• 






}0Ê_^nm><"' »./ ' 






5B 



m 




5C 



5D 



5E 



FlG 5. Spatola. A: Pipistrellus pygmaeus. B: Nyctalus lasiopterus. C: Vespertilio murinus. D: 
Hypsugo savii. E: Myotis myotis. 



388 R- PIERALLINI ET AL. 

3 Sulla parte centrale del fusto le scaglie sono caliciformi (Fig. 6A). 

I calici possono essere presenti anche sulla base del fusto . . Pipistrellus kuhlfi 

Sulla base e sulla parte centrale del fusto la maggior parte delle scaglie 

non è caliciforme, bensì a forma di imbuto (Fig. 6B, C) 4 

4 Sulla spatola le scaglie sono divergenti e spaziate (Fig. 6D). Sulla parte 

apicale del fusto le scaglie sono talvolta caliciformi 5 

Sulla spatola le scaglie sono divergenti, ma più fitte e appiattite in avanti 

(Fig. 6E). Non vi sono spazi tra le scaglie. Di norma mancano le scaglie 
caliciformi (difficile) Pipistrellus nathusii 

5 Di norma le scaglie sono meno divergenti (Fig. 6C). Il fusto è più esile 

(difficile) Nyctalus leislerf 

Di norma le scaglie sono più divergenti (Fig. 6B). Il fusto è più robusto, 

con un numero maggiore di strutture caliciformi 
Pipistrellus pipistrellus, P. pygmaeus^ 

6 Sulla parte apicale del fusto le scaglie formano una struttura "a perle" 
(cfr. Fig. 7F). Sulla sua base sono invece alternate, robuste e angolose. 
La radice è corta e può presentare un rigonfiamento. Pelo marrone 

Vespertilio murinus 

Sulla parte apicale del fusto le scaglie non formano una struttura "a per- 
le" (Fig. 6F). Il fusto è più esile. La radice è lunga e senza rigonfiamenti. 

Pelo marrone chiaro Nyctalus noctula, N. lasiopterus 9 

1 Le scaglie del fusto hanno una struttura molto asimmetrica e formano 

una caratteristica linea ondulata marcata e regolare (Fig. 7A) 

Miniopterus schreibersi 

L'ondulazione del fusto è poco marcata, irregolare oppure assente (Fig. 

7B, C, D, E, F) 8 

8 Lungo tutto il fusto le scaglie sono appiattite e poco strutturate (Fig. 7E). 

L'ondulazione del fusto è pressoché assente 9 

Sulla base e sulla parte centrale del fusto le scaglie sono sempre ben 
strutturate, anche se manca ondulazione. 

Sulla parte apicale le scaglie possono invece anche essere poco strutturate . . 10 

9 La radice del pelo è sempre lunga e diritta (Fig. 8A) . Sulla parte apicale 
del fusto le scaglie sono simmetriche e poco strutturate (cfr. Fig.7E) 

Eptesicus serotinus 1 ® 

La radice del pelo è più corta e di norma curva o con rigonfiamento (Fig. 

8B). Sulla parte apicale del fusto le scaglie formano una struttura "a 
perle" (cfr. Fig. 7F) Barbastella barbastellus 



6 Per le specie del genere Pipistrellus si rimanda il lettore al lavoro di Keller (1986). 

7 Lo sterco di N. leisleri (d > 2 mm) è più grande di quello di P. pipistrellus/ pygmaeus (1 < 8 
mm, d < 2 mm) (1 = lunghezza; d = diametro). 

8 Di norma in P. pipistì-ellus le scaglie della spatola formano un angolo con l'asse principale 
prossimo ai 45°, mentre in P. pygmaeus l'angolo è inferiore ai 45°. Questo criterio è però da 
considerarsi difficile e incerto. 

9 Lo sterco di N. noctula (1 > 1 cm; d ~ 4 mm) è più grande di quello di V. murinus (d < 4 mm). 
10 Lo sterco di E. serotinus (1 ~ 1 cm; d ~ 3 mm) è più grande di quello di B. barbastellus. 



CHIAVE DI DETERMINAZIONE DEI CHIROTTERI 



389 



**#*#######*• 



6A 



0-0mm-***+* 



6B 



4Ê -éÊ 4 ~*Ë A ^^. 



30 m m 



A «ni 






■•.. •». 



6C 



„ j 



6D 




*•• 






50 pm 



6E 



6F 



Fig. 6. A: Pipistrellus kuhli, parte centrale del fusto. B: Pipistrellus pygmaeus, parte centrale del 
fusto. C: Nyctalus leisleri, parte centrale del fusto. D: Pipistrellus pipistrellus, spatola. E: 
Pipistrellus nathusii, spatola. F: Nyctalus noctula, parte apicale del fusto. (Ingrandimento: D, E, 
1000 volte; A, B, C, F, 400 volte). 



10 Sulla parte apicale del fusto le scaglie formano per un lungo tratto una 
struttura "a perle" (Fig. 7F). Sulle parti centrale e apicale del fusto 
l'ondulazione è quasi assente Hypsugo savii, Eptesicus nilssoni 



390 



R. PIERALLINI ET AL. 




7E 




Fig. 7. Fusto A: Miniopterus schreibersi. B: Myotis daubentoni. C: Myotis myotis. D: Plecotus 
auritits. E: Eptesicus serotinus. F: Hypsugo savii, parte apicale del fusto. 



Sulla parte apicale del fusto la struttura "a perle" è presente solo per un 
breve tratto oppure è assente. In alcuni tratti il fusto è chiaramente on- 
dulato (Fig. 7B, C. D) 11 

11 Radice corta (< 110 nm) con scaglie corte e subito divergenti (Fig. 8C) 

Plecotus spp. 

Radice lunga (> 110 nm) con scaglie lunghe che divergono gradual- 
mente (fig. 8D, E), oppure con rigonfiamento (Fig. 8F) 12 

12 La radice presenta sovente un rigonfiamento (Fig. 8F) .... Myotis emarginatus 
La radice è stretta, senza rigonfiamento (Fig. 8D, E) Myotis spp. 11 



11 Lo sterco di M. myotis e M. blythi si distingue da quello delle altre specie di Myotis per le 
maggiori dimensioni (1 > 1 cm; d ~ 4 mm). 



CHIAVE DI DETERMINAZIONE DEI CHIROTTERI 



391 




8A 



8B 



£-t% 



*^&mt&imm*i!i!8mfàto 



■4 *. 



•.,,,, ,,.:" /-•" '"" ' 




Fio. 8 
Radice. A: Eptesicus serotinus. B: Barbastello, barbastellus. C: Plecotus auritus. D: Myotis 
mystacinus. E: Myotis myotis F: Myotis emarginatus. 

DISCUSSIONE 

Benedict (1957), che è stato uno dei pionieri in questo campo, aveva già osser- 
vato una considerevole variabilità di strutture nei peli dei Microchirotteri. Il nostro 
lavoro conferma le sue asserzioni, e rivela che molte specie e quasi tutti i generi 
possiedono una morfologia del pelo tipica. Per esempio Tadarida teniotis è l'unica 
specie a presentare una vera struttura coronale (già citata in Tupinier 1973), mentre 
Miniopterus schreibersi si distingue per una struttura asimmetrica particolare. Il genere 
Pipistrellus ha pure caratteristiche peculiari, tali da permettere l'identificazione delle 
singole specie (Keller, 1986). Lo stesso vale per i Rinolofidi. Per i Myotis la situazione 
è invece diversa: l'identificazione del genere non è un problema, ma le diverse specie 



392 R. PIERALLINI ETAL. 

sono assai simili tra loro e si riesce a separare unicamente M. emarginatus (nel caso di 
peli prelevati dallo sterco, considerato la maggiore taglia, si arriva a separare anche M. 
my otis e blythi dalle altre specie). 

Pregio della chiave qui presentata è quello di permettere l'identificazione degli 
animali, indipendentemente dal loro sesso e dalla loro età (Keller & Moeschler, 1988), 
unicamente sulla base dei peli contenuti nello sterco raccolto nei rifugi, limitando così 
significativamente il disturbo causato agli animali. Le informazioni fornite da questo 
tipo di analisi rappresentano un importante aiuto per decidere se e dove proseguire le 
indagini con le catture per avere ulteriori informazioni sulla colonia (conferma della 
specie, riproduzione, età, sesso). Nei rifugi abbandonati inoltre si possono ottenere 
informazioni sulla distribuzione passata di specie in forte regresso o estinte (p. es. 
Rhinolophidae). 

RINGRAZIAMENTI 

Ad eccezione della parte dedicata ai Rinolofidi, eseguita al Museo di storia 
naturale di Ginevra, lo studio è stato interamente finanziato dal Museo cantonale di 
storia naturale di Lugano (Svizzera), che ha anche fornito il supporto logistico e 
l'attrezzatura necessaria a realizzare le osservazioni e le foto (microscopio e tele- 
camera). Ringraziamo in particolare il suo direttore, Filippo Rampazzi, e il conser- 
vatore del settore Vertebrati, dr. Alessandro Fossati, che hanno creduto nella sua 
realizzazione. Si ringraziano i Musei di storia naturale di Ginevra, Lugano, Coirà, 
Barcellona, la dr. Marianne Haffner del Museo zoologico dell'Università di Zurigo, il 
CPT e tutti coloro che hanno fornito i campioni di peli. Grazie anche a Neria Roemer 
per l'assistenza neh' utilizzo della fotocamera. Infine si ringrazia dr. A. Fossati, M. 
Moretti, Marzia Roesli e F. Rampazzi per la lettura dell'articolo e i suggerimenti. 

BIBLIOGRAFIA 

Benedict, FA. 1957. Hair structure as a generic character in bats. University California 

Publications Zoology 59: 285-548. 
Charvet, C. & Keller, A. 1989. Une méthode douce d'identification des mammifères: la struc- 
ture fine des poils. Le Rhinolophe 6:19-26. 
Debrot, S., FrvAZ, G., Mermod, C. & Weber, J.-M. 1982. Atlas des poils de mammifères 

d'Europe. Institut Zoologie Université Neuchâtel, 208 pp. 
Dove, C.J. & Peurach, S.C. 2001. The use of microscopie hair characters to aid in identification 

of bat involved in a damaging aircraft strike. Bat Research News 42(1): 10-11. 
Dulic, B. 1978. Morphology of the hair of Pipistrellus savii Bonaparte 1837. Proceedings 4th 

International bat research conference. Kenya literature, 51-61 pp. 
Hausser, J. 1995. Mammiferi della Svizzera. Birkhäuser Verlag, Basel-Boston-Berlin, 501 pp. 
Jones, G. & Barrati, E.M. 1999. Vespertilio pipistrellus Schreber, 1774 and V. pygmaeus 

Leach, 1825 (currently Pipistrellus pipistrellus and P. pygmaeus; Mammalia, Chiro- 

ptera): proposed designation of neotypes. Bulletin of Zoological Nomenclature 56: 182- 

186. 
Keller, A. 1978. Détermination des mammifères de la Suisse par leur pelage: I. Talpidae et 

Soricidae. Revue suisse de Zoologie 85(4): 758-761. 
Keller, A. 1980. Détermination des mammifères de la Suisse par leur pelage: II. Diagnose des 

familles. III. Lagomorpha, Rodentia (partim). Revue suisse de Zoologie 87(3): 781-796. 



CHIAVE DI DETERMINAZIONE DEI CHIROTTERI 393 



Keller, A. 1986. Etude comparative de la structure fine des poils des Pipistrelles d'Europe 
(Mammalia : Chiroptera). Revue suisse de Zoologie 93(2): 409-415. 

Keller, A. & Moeschler, P. 1988. Résolution d'un problème d'identification d'une jeune 
Pipistrellus kuhli (Natterer) in Kuhl, par analyse de la structure fine des poils. Le 
Rhinolophe 5: 25-30. 

Kiefer, A. & Veith, M. 2001 . A new species of long-eared bat from Europe (Chiroptera: Vesper- 
tilionidae). Myotis 39: 5-16. 

Meyer, W., Seger, H. & Hülmann, G. 1995. Remarks on Specific Adaptive Scale Structure of 
the Hair Cuticle in Some European Bats. European Journal of Morphology 33(5): 
509-513. 

Meyer, W., Seger, H., Hülmann, G. & Neurad, K. 1997. A computer-assisted method for the 
determination of hair cuticula patterns in mammals. Berliner und Münchener Tier- 
ärztliche Wochenschrift 110: 81-85. 

Teerink, B.J. 1991. Hair of West European Mammals. Cambridge University Press, 224 pp. 

Tupinier, Y. 1973. Morphologie des poils de Chiroptères d'Europe occidentale par étude au 
microscope électronique à balayage. Revue suisse de Zoologie 80(2): 635-653. 



Revue suisse de Zoologie 111 (2): 395-424; juin 2004 



Taxonomie der Bernstein- Waldschabe Ectobius vittiventris 
(A. Costa, 1847) (Blattodea: Blattellidae) und ihre Verbreitung 
in der Schweiz 

Hannes BAUR 1 , Isabelle LANDAU LÜSCHER 2 , Gabi MÜLLER 2 , 
Marcus SCHMIDT 2 & Armin CORAY 3 

1 Abteilung Wirbellose Tiere, Naturhistorisches Museum, Bernastrasse 15, 
CH-3005 Bern, Schweiz. E-Mail: hannes.baur@nmbe.unibe.ch (Korrespondenz) 

2 Beratungsstelle Schädlingsbekämpfung (BSB), Umwelt- und Gesundheitsschutz 
Zürich (UGZ), Walchestrasse 33, CH-8035 Zürich, Schweiz. 

3 Naturhistorisches Museum, Augustinergasse 2, CH-4001 Basel, Schweiz. 



Taxonomy of the field-dwelling cockroach Ectobius vittiventris (A. Costa, 
1847) (Blattodea: Blattellidae) and its distribution in Switzerland. - In 

this paper we examine the taxonomy of E. vittiventris (Ectobiinae) and its 
distribution and phenology in Switzerland. An illustrated key and a diagnosis 
are provided to separate the adults from synanthropic cockroaches and from 
some native Ectobius spp. Descriptive notes are included for nymphs and 
oothecae. We furthermore discuss the taxonomic status of the species and 
provide information on the type locality and type series, and the same is 
given for its junior synonyms. In Switzerland E. vittiventris is found on both 
sides of the Alps. In the northern part it is mainly confined to urban areas of 
the central plateau. On the southern slopes of the Alps it occurs in the can- 
tons Ticino, Valais and Grisons. Our data indicate a rather recent introduction 
and colonization of urban areas in northern Switzerland, where the species 
has been recorded mostly after 1985. The altitudinal distribution generally 
ranges from 260 to about 700 m, with very few records up to 1400 m in the 
south. Adults are encountered from May to December, with a peak frequency 
in August. The presence of very small nymphs in summer and of larger 
nymphs in winter suggests a two-year life-cycle. E. vittiventris is frequently 
encountered inside houses, but apparently it cannot survive under such con- 
ditions. A special treatment or control is thus unnecessary. Finally, E. vitti- 
ventris is recorded for the first time from Germany (Baden- Württemberg). 

Key-words: Blattodea - morphology - identification - phenology - pests - 
Blattella - Blatta - Ectobius - Periplaneta - Supella - Switzerland. 

EINLEITUNG 

Mit weltweit nur etwa 4000 Arten (Roth, 1991) sind die Schaben oder Blattodea 
relativ artenarm. Ihren grössten Artenreichtum weist die ursprüngliche Insektengruppe 



Manuskript angenommen am 16.01.2004 



396 H. BAURET AL. 

in den Tropen und Subtropen auf (Beier, 1967; Roth, 1991). In der Schweiz kommen, 
je nach Quelle, nur gerade 12-15 Arten vor (Fruhstorfer, 1921; Harz, 1957b; Princis, 
1971; Harz & Kaltenbach, 1976). Davon gehören 8-10 Arten zu den frei lebenden 
Wald- und Kleinschaben der Gattungen Ectobius Stephens und Phyllodromica Fieber 
(Blattellidae: Ectobiinae), welche in faunistischer Hinsicht jedoch sehr ungenügend 
erforscht sind. Wald- und Kleinschaben sind scheue und wenig auffällige Bewohner 
von Hecken und Waldrändern (Harz, 1957b). Viele Arten leben versteckt im Laub oder 
unter Steinen. Bei der Eiablage werden die Eier mit einem Drüsensekret umgeben, 
welches zu einem festen Paket, der sogenannten Oothek, erhärtet (Harz, 1960; Beier, 
1967). Die Oothek wird vom Weibchen zuweilen noch mehrere Tage am Abdomen- 
ende herumgetragen (Harz, 1960) und schliesslich in die Laubstreu abgelegt (Brown, 
1973c) oder vergraben (Harz, 1957a). Für einige Arten wurde ein zweijähriger Ent- 
wicklungszyklus festgestellt (Morvan, 1972; Brown, 1973c, 1980; Holusa & Kocärek, 
2000). E. lapponicus und E. pallidus (Olivier) können sich fakultativ parthenogene- 
tisch vermehren (Brown, 1973b). Für die mitteleuropäischen Waldschaben werden 
Brachygaster minuta (Olivier) (Hymenoptera: Evaniidae) als Eiparasitoid (Brown, 
1973a) sowie Rhipidius quadriceps Abeille de Petrin (Coleoptera: Rhipiphoridae) als 
Nymphalparasitoid (Besuchet, 1956) erwähnt. Wald- und Kleinschaben ernähren sich 
vor allem von verschiedenen pflanzlichen Stoffen (Harz, 1960), allerdings fehlen 
gezielte Freilanduntersuchungen zu den einzelnen Arten. 

Neben den freilebenden Schaben treten mehrere eingeschleppte, ursprünglich 
aus südlichen Gebieten stammende Arten regelmässig im Bereich von menschlichen 
Siedlungen auf. In Mitteleuropa gehören zu diesen synanthropen, kosmopolitischen 
Arten in erster Linie die Deutsche Schabe Blattella germanica, die Braunband-Schabe 
Supella longipalpa, die Küchenschabe Blatta orientalis, sowie zwei Arten der Gattung 
Periplaneta (vgl. Weidner, 1993). Durch Massenvermehrung können sie ausser- 
ordentlich schädlich werden und stellen als potenzielle Krankheitsüberträger an expo- 
nierten Orten (z. B. in Spitälern) ein Gesundheitsrisiko dar (Beier, 1967; Mallis, 1997). 
Wo sie auftreten, müssen sie daher konsequent bekämpft werden. 

Im Laufe der letzten 10-15 Jahre wurden in der Nordschweiz verschiedentlich 
Meldungen über das Auftreten von Waldschaben in Wohnungen bekannt (Herger, 
2000; Landau Lüscher et al., 2003). Ausgehend von den deutsch-schweizerischen 
Zentren wie Basel, Bern und Zürich wurde man auf die Tiere zunehmend auch in der 
französischen Schweiz sowie auf dem Land in kleineren Städten aufmerksam. Zu- 
nächst wurde angenommen, es handle sich bei diesen ungewöhnlichen Funden um die 
Lappland-Waldschabe E. lapponicus (vgl. Landau et al, 1999), welche in Nordeuropa 
gelegentlich schon früher in Häusern festgestellt wurde (Harz, 1957b; Weidner, 1972; 
Abraham, 1979). In einer eingehenden Untersuchung konnten die betreffenden 
Exemplare jedoch eindeutig als Bernstein-Waldschaben E. vittiventris bestimmt 
werden (Baur, Bohn, Coray, unpubl.). Obwohl sich die Art in Häusern nicht vermehren 
kann und keinen Schaden anrichtet (Landau Lüscher et al., 2003), ist das Auftreten für 
die betroffenen Personen dennoch oft unangenehm. Die Bernstein-Waldschabe gleicht 
bei oberflächlicher Betrachtung nämlich stark der Deutschen Schabe, welche in 
Wohnungen schädlich wird und deshalb eine Behandlung durch eine Schädlings- 
bekämpfungsfirma erforderlich macht (Pospischil, 1996). 



ECTOBIÜS VITTIVENTRIS (BLATTODEA) 397 

Die Bemstein-Waldschabe ist eine südeuropäische Art (Ramme, 1951; Harz, 
1957b; Harz & Kaltenbach, 1976), die in der Schweiz die Nordgrenze ihrer Aus- 
breitung erreicht. Bisher war sie nur von der Alpensüdseite, z. B. dem Kanton Tessin, 
bekannt (Fruhstorfer, 1921, sub E. neolividus). Das scheinbar plötzliche Auftreten in 
der Nordschweiz, noch dazu in Wohnungen, stellte deshalb eine grosse Überraschung 
dar. In der vorliegenden Arbeit werden nun erstmals Daten zur Taxonomie der Bern- 
stein-Waldschabe und ihrer Verbreitung in der Schweiz ausgewertet. Während 
mehreren Jahren haben wir alle verfügbaren Angaben zur Verbreitung gesammelt. Von 
besonderer Bedeutung waren dabei die Erhebungen, welche seit Ende der Achtziger- 
jahre vom Umwelt- und Gesundheitsschutz Zürich (UGZ) durchgeführt wurden. 
Ergänzend wurde das Material verschiedener Museen in der Schweiz und im Ausland 
durchgesehen. Neben der horizontalen sowie vertikalen Verbreitung interessierte uns 
insbesondere die Phänologie der einzelnen Stadien, um indirekt Hinweise auf den 
Entwicklungszyklus der Art zu erhalten. Direkte Beobachtungen hierzu fehlten bisher 
völlig. Ferner haben wir die wichtigsten Merkmale an zahlreichen Museumsbelegen 
untersucht und in einer Diagnose zusammengestellt. Ein illustrierter Bestimmungs- 
schlüssel sollte es schliesslich auch dem NichtSpezialisten erlauben, die Bernstein- 
Waldschabe sicher zu erkennen. 

MATERIAL UND METHODEN 

Für die vorliegende Studie wurden insgesamt 2000 Datensätze (siehe unten) aus 
der Schweiz (1984) und Teilen von Deutschland (4) und Italien (12) ausgewertet. Die 
Untersuchungen zur Morphologie, Verbreitung, Höhen Verbreitung und Phänologie 
basieren ausschliesslich auf Daten aus der Schweiz. Grosse Unterschiede hinsichtlich 
ihrer Qualität machten eine Unterteilung dieser Daten in zwei Gruppen notwendig, 
welche in den Analysen jeweils getrennt behandelt wurden. Die eine Gruppe wurde 
von der Beratungsstelle Schädlingsbekämpfung des Umwelt- und Gesundheitsschutzes 
Zürich (UGZ) von 1991 bis 1999 und 2001 zusammengetragen und wird als UGZ- 
Daten bezeichnet (1637 Datensätze, vgl. Tab. 1, Appendix 1). Seit 1991 wurden an der 
Beratungsstelle alle Schädlingsmeldungen systematisch auf EDV erfasst, wobei aber 
nur die Zahl der Meldungen pro Art und Fundort aufgenommen wurde. Die Anzahl der 
Individuen, deren Stadien (z. B. Nymphen oder Imagines) etc. wurde dagegen nicht 
systematisch erhoben. Belege aus diesen Jahren fehlen weitgehend. Seit Beginn der 
Datenerfassung konnte nun eine Art der Gattung Ectobius festgestellt werden, die zwar 
leicht als Waldschabe zu erkennen war, deren Artzugehörigkeit aber lange Zeit unklar 
blieb. Erst eine Bestimmung aller Individuen des Jahres 2000 machte deutlich, dass es 
sich fast ausschliesslich um E. vittiventris handelte (Anteil anderer Ectobius-Arten < 
0,5%). Das gleiche Resultat ergaben die wenigen erhaltenen Belege aus den Jahren 
1999 und 2001 (vgl. Appendix 2). Bei der Auswertung der UGZ-Daten konnte deshalb 
davon ausgegangen werden, dass es sich bei den nicht mehr überprüfbaren Wald- 
schaben der anderen Jahre höchstwahrscheinlich ebenfalls um E. vittiventris handelt. 
Die UGZ-Daten wurden aber wie erwähnt jeweils getrennt analysiert, damit die 
Grundlage der Resultate überall nachvollziehbar bleibt. Demgegenüber bildete das 
UGZ-Material des Jahres 2000, welches vollumfänglich im Naturhistorischen Museum 
Bern aufbewahrt wird, zusammen mit allem anderen Museumsmaterial die Basis der 



398 H. BAURET AL. 



Tabelle 1: UGZ-Daten 1991-2001. Angegeben ist die Anzahl Datensätze von Ectobius 
vittiventris pro Jahr und Monat (vgl. Material und Methoden; für das Jahr 2000 sind die ein- 
zelnen Exemplare in Appendix 2 aufgelistet). "Total" zeigt die Gesamtzahl der Datensätze 
pro Jahr. "% Ev" entspricht dem Anteil (in %) von E. vittiventris an der Gesamtheit aller 
Tiermeldungen pro Jahr (nur für 1991-2000 berechnet). *Durch eine öffentliche Veranstaltungen 
des UGZ zum Thema "Waldschaben" kamen im Oktober 1999 in Wirklichkeit 100 Meldungen 
zustande. Diese aussergewöhnlich hohe Zahl stellt aber eindeutig einen Artefakt dar, da die 
Bevölkerung speziell dazu angeregt wurde, Funde von Waldschaben dem UGZ zu melden. Für 
die Auswertung (Abb. 22, 24) wurde deshalb der gerundete Oktober-Mittelwert der anderen 
Jahre (= 16) verwendet. 



Jahr 


Jan 


Feb 


Mär 


Apr 


Mai 


Jun 


Jul 


Aug 


Sep 


Okt 


Nov 


Dez 


Total 


%Ev 


1991 

















1 


5 


15 


6 


3 





3 


33 


2.3 


1992 








2 





1 


6 


35 


54 


4 


6 


1 





109 


5.9 


1993 














1 


4 


8 


19 


12 


12 


9 


1 


66 


3.3 


1994 





2 








1 


4 


56 


41 


20 


20 


14 


3 


161 


8.0 


1995 














1 


4 


37 


91 


27 


22 


19 


4 


205 


7.7 


1996 


2 


3 








1 


21 


61 


77 


38 


25 


5 


1 


234 


10.2 


1997 





1 


1 





7 


8 


32 


56 


24 


15 


3 


2 


149 


7.2 


1998 





1 





2 


2 


28 


103 


91 


28 


14 


7 


1 


277 


11.5 


1999 


4 





2 


1 


5 


8 


18 


41 


32 


*16 


34 


4 


165 


6.5 


2000 


5 


4 


3 


5 


12 


36 


49 


75 


24 


13 


13 


2 


241 


10.2 


2001 


1 


1 


3 


3 


9 


11 


48 


101 


25 


25 


9 


2 


238 


- 


1991-2001 


12 


12 


11 


11 


40 


131 


452 


661 


240 


171 


114 


23 


1878 


100 



MUS-Daten (347 Datensätze von 600 Exemplaren und 31 Ootheken, vgl. Appendix 
2). Für diese Daten wurde jedes Exemplar bestimmt und getrennt nach Geschlecht oder 
Stadium (Imagines oder Nymphen) erfasst. Die genauen Fundumstände wurden 
systematisch notiert. Die MUS-Daten bildeten die bevorzugte Grundlage für die 
meisten unserer Untersuchungen, z. B. zur Morphologie, Verbreitung und Phänologie. 
Messungen wurden an Alkoholmaterial, unter einem Stereomikroskop bei 8- 
(Gesamt-, Körper- und Deckflügellänge), 30- (Pronotumlänge) bzw. 20-f acher (Oothek) 
Vergrösserung, mit Hilfe eines geeichten Okular-Mikrometers (12 mm unterteilt in 120 
Einheiten), durchgeführt, anschliessend wurden die Werte in Millimeter umgerechnet. 
Die betreffenden Exemplare wurden mit einer Etikette mit dem Vermerk "Baur" und 
einer Seriennummer versehen. Die Gesamtlänge entsprach der Distanz vom Kopf bis 
zur Spitze der Deckflügel, die Körperlänge derjenigen vom Kopf bis zum Apex der 
Subgenitalplatte. Der linke Deckflügel wurde in situ von der Basis bis zur Spitze 
gemessen, die Pronotumlänge entlang der zentralen Längsachse. Die Auswahl der 
Individuen (vgl. Appendix 2) erfolgte zufällig. In der morphologischen Terminologie 
folgen wir Roth (1991), einige Begriffe werden zusätzlich in den Abb. 6-8 erläutert. 
Quantitative Angaben beziehen sich auf die Gesamtheit des MUS-Materials (nur 
Schweizerdaten) und sind wie folgt definiert: sehr selten < 1%, selten 1-10%, gele- 
gentlich > 10-25%, manchmal > 25-50%, oft > 50-75%, häufig > 75-90%, meist > 90- 
99%, normalerweise > 99% des vorliegenden Materials. Für die Auswertung der 
Messreihen wurde SPSS 11.00 (2001) verwendet, für die Erstellung der Verbreitungs- 
karte DMAP, Version 6.5 (1998). 



ECTOBIUS VITTIVENTRIS (BLATTODEA) 



399 



Abkürzungen der Kantone der Schweiz 



AG 

BE 

BL 

BS 

GE 

GR 

JU 

LU 

NE 

NW 



Aargau 

Bern 

Basel Land 

Basel Stadt 

Genf 

Graubünden 

Jura 

Luzern 

Neuenburg 

Nidwaiden 



ow 


Obwalden 


SG 


St. Gallen 


SH 


Schaffhausen 


SO 


Solothurn 


SZ 


Schwyz 


TI 


Tessin 


VD 


Waadt 


VS 


Wallis 


ZG 


Zug 


ZH 


Zürich 



Abkürzungen der Sammlungen 

CAN Coll. Adolf Nadig, in MHNG 

CGA Coli. Georg Artmann, Ölten, Schweiz 

CKH Coli. Kurt Harz, in MHNG 

ETHZ Eidgenössische Technische Hochschule, Zürich, Schweiz 

MHNG Muséum d'histoire naturelle, Genève, Schweiz 

NHMB Naturhistorisches Museum, Basel, Schweiz 

NMBE Naturhistorisches Museum, Bern, Schweiz 

NML Natur-Museum, Luzern, Schweiz 

UGZ Beratungsstelle Schädlingsbekämpfung, Umwelt- und Gesundheitsschutz, 

Zürich, Schweiz, in NMBE 
ZMB Zoologisches Museum, Humboldt Universität, Berlin, Deutschland 



RESULTATE 

Ectobius vittiventris (A. Costa) - Bernstein- Waldschabe 

Blatta vittiventris A. Costa, 1847: 111-112. Syntypen 6 6 99, Italien, Campania: Monte 
Matese, Cusano, coll. ? [Nicht untersucht]. 

Ectobius neolividus Fruhstorfer, 1921: 78-79. Syntypen 6 699, Schweiz: Graubünden, 
Südtessin, coll. CAN, ZMB [\ 6,2 9 untersucht, s. Appendix 2]. Syn. Ramme (1923: 
130). 

Ectobius grandis Ramme, 1922: 183-184. Syntypen 10 6, 41 $,6 Nymphen, Norditalien: 
Gardasee, Trient, Val Sugana, Atzwang (nahe Bolzano), coll. ? [Nicht untersucht. Nach 
Auskunft von Frau Isolde Dorandt, ZMB, befinden sich keine Exemplare, die eindeutig 
als Syntypen von E. grandis zu erkennen wären, in der coli. Ramme; weitere 
Abklärungen vor Ort und in anderen Sammlungen wären daher wünschenswert]. Syn. 
Ramme (1923: 130). 

Ectobius vitreus Ramme, 1923: 134; Taf. 2, Abb. 8. Syntypen 8 6,5 9, Italien, Kroatien, 
Rumänien, coli. ZMB [1 6 , 2 9 untersucht, s. Appendix 2]. Syn. Ramme (1951: 42). 

Status 

Aufgrund der heutigen Auffassung (z. B. Harz & Kaltenbach, 1976) sind die 
obigen nominellen Taxa als Synonyme von E. vittiventris zu betrachten. Nach Bohn (in 
litt.) ist es jedoch möglich, dass sich unter diesem Namen mehrere Arten verbergen und 
die Populationen nördlich vom Apennin (Italien) nicht derselben Art angehören wie die 
Populationen südlich davon. Die Klärung des genauen Status aller Populationen muss 
freilich einer umfassenderen Studie vorbehalten bleiben und war nicht das Ziel der 



400 H. BAURET AL. 

vorliegenden Arbeit. Das im Appendix aufgelistete Material aus der Schweiz, aus 
Deutschland, Nord- und Mittelitalien gehört unseres Erachtens aber nur einer Art an. 

Diagnose 

Imagines: Eine der grössten europäischen Ectobius-Aiten, Länge: Gesamt S 
12.4-14.6 mm, 9 9.3-11.9 mm, Körper S 10.3-12.3 mm, 9 8.4-11.0 mm, Pronotum 
6 2.2-2.6 mm, 9 2.3-2.8 mm, Deckflügel S 10.6-12.1 mm, 9 7.8-10.0 mm (vgl. Tab. 
2). S von oben betrachtet deutlich schlanker als das 2 . Körper überwiegend ocker- 
farben (Abb. 15-16). Kopf gelblich bis orangebraun, Gesicht manchmal dunkler. 
Discus des Pronotums ± einfarbig bernsteinfarben, gelegentlich mit unauffälligen, 
dunklen Flecken (Abb. 2, 15, 16); Saum heller als Discus, ± durchsichtig. Flügel über- 
ragen das Abdomen beim S um ca. 2.0-3.1 mm, beim 9 um ca. 0.3-2.5 mm; â mit 
deutlich längeren Deckflügeln als die 9 , Index Deckflügel- / Pronotumlänge â 4.5- 
4.9, 2 3.1-3.8 (vgl. Tab. 2); Farbe der Deckflügel einfarbig glasig gelblich, fein 
gemustert, aber ohne grössere dunkle Flecken (Abb. 4). Hinterflügel leicht gebräunt, in 
Ruhelage beim 9 ungefähr so lang wie die Deckflügel, beim â normalerweise ca. 0.5 
mm länger, Analfeld der Hinterflügel mit spiralig aufgerolltem Apikaidreieck. Beine 
gelblich, Hüften und Schenkel manchmal recht stark gebräunt; Fussglieder meist mit 
schmalem braunen Ring am Apex. Unterseite der Schenkel nur mit vereinzelten 
Dornen (Enddornen nicht mitgerechnet): Vorderschenkel l-3(-4), Mittelschenkel (l-)2- 
4, Hinterschenkel l-3(-5), die Bedornung an Vorder- und Hinterschenkel meist nur in 
der proximalen Hälfte. Abdomenoberseite braun, am Rand und an der Spitze oft auf- 
gehellt (Abb. 7). Drüsengrube queroval, nur an den Rändern mit Haaren besetzt, in der 
Mitte kahl und höchstens mit schwacher Längserhebung (Abb. 7, 10-12). Zeichnung 
der Abdomenunterseite hell- bis schwarzbraun; in der distalen Hälfte drei dunkle 
Längsstreifen erkennbar, diese bei dunklen Exemplaren oft ineinander verschmolzen. 
Subgenitalplatte des â im Umriss dreieckig, mit nur einem Stylus an der Spitze, 
höchstens bis zur Mitte von der Supraanalplatte bedeckt (Abb. 7). Cerci schlank, ± 
gelblich, leicht angedunkelt (Abb. 7). Stylus des â relativ gross, gelblich, oberseits mit 
honigfarbenem, dichtem Haarstreifen (Abb. 7). Linker Phallomer mit relativ 
schlankem Haken (Abb. 14). 

Tabelle 2: Masse verschiedener Merkmale von Ectobius vittiventris. 

Merkmal Minimum Maximum Mittelwert Standardfehler n 

Gesamtlänge (mm) 

Körperlänge (mm) 

Pronotumlänge (mm) 

Deckflügellänge (mm) 

Deckflügel / Pronotum 



ö 


12.4 


14.6 


13.59 


0.0835 


30 


9 


9.3 


11.9 


10.68 


0.0939 


30 


d 1 


10.3 


12.3 


11.20 


0.0910 


30 


9 


8.4 


11.0 


9.34 


0.1156 


30 


d 1 


2.2 


2.6 


2.44 


0.0149 


30 


9 


2.3 


2.8 


2.53 


0.0208 


30 


d 


10.6 


12.1 


11.38 


0.0675 


30 


9 


7.8 


10.0 


8.85 


0.1004 


30 


d 


4.5 


4.9 


4.66 


0.0193 


30 


9 


3.1 


3.8 


3.50 


0.0246 


30 



ECTOBIUS VITTIVENTRIS (BLATTODEA) 



401 




L_ 














Abb. 1-5 

Pronotum <5: (1) Blattella germanica von Bern/BE, (2) Ectobius vittiventris von Riehen/BS, (3) 
£. lucidus von Aesch/BL. Linker Deckflügel 9 : (4) E. vittiventris von Bottmingen/BL, (5) E. 
lapponicus von Basel/BS. Massstab 1 mm. Zeichnungen A. Coray. 



402 H. BAURET AL. 

Nymphen: Die einzelnen Nymphenstadien wurden nicht getrennt untersucht. 
Körperlänge ca. 2.3-7.0 mm. Färbung und Zeichnung ähnlich den Imagines und 
dadurch leicht von den synanthropen Arten und dunklen Ectobius- Asten wie E. 
sylvestris zu unterscheiden: Thorax gelblich, Discus des Pronotums einfarbig bern- 
steinfarben, rundlich; Pronotum und Mesothorax oft fein gepunktet. Flügelansätze 
gelblich. Abdomen braun, oft hell gezeichnet. Inwieweit sich die Nymphen von denen 
weiterer Ectobius-Aiten unterscheiden, konnte nicht untersucht werden. 

Oothek: Länge 2.9-4.9 mm, Breite in der Mitte 2.1-2.7 mm (n=15). Braun bis 
schwarzbraun. Äussere Form der Oothek (Abb. 9) in Längsrichtung leicht gekrümmt. 
Trennwände der Eikammern deutlich als feine Querlinien durchscheinend. Oberfläche 
schwach skulpturiert, auf der ganzen Breite mit zahlreichen, feinsten Längsrippen. 
Oberkante mit ca. 15-25 Zähnchen; Unterseite mit breiter und recht tiefer Längsfurche. 
Oothek mit 12-23 Eikammern (n=15). 

Bestimmungsschlüssel 

Mit dem folgenden Bestimmungsschlüssel lassen sich die Imagines von synan- 
thropen Schaben sowie von jenen Freilandarten der Gattung Ectobius, welche 
vereinzelt auch in Häusern angetroffen wurden, unterscheiden. Für die Bestimmung 
der übrigen Wald- und Kleinschaben verweisen wir auf die Arbeiten von Harz (1957b), 
Harz & Kaltenbach (1976) und Bohn (2000). 

1 Kleinere Arten, Gesamtlänge ca. 8-15 mm 2 

Grössere Arten, Gesamtlänge ca. 1 8-40 mm 7 

2 Deckflügel relativ hell, glasig gelblich bis hellbraun, gelegentlich mit 
einigen dunklen Flecken (Abb. 4, 5) (Achtung: das dunkle Abdomen 
kann bei gewissen Arten durchschimmern und einen falschen Eindruck 

der Deckflügelfarbe erwecken) 3 

Deckflügel mit 1-2 breiten braunen Querbinden, aber nie mit dunklen 

Flecken Supella longipalpa (Fabricius) - Braunband-Schabe 

[Discus des Pronotums einfarbig braun, rundlich. Flügel überragen beim S das 
Abdomenende um 2-3 mm, beim 9 lassen sie es frei. Hinterflügel in Ruhelage 
kürzer als Deckflügel, Analfeld der Hinterflügel mit gefaltetem Apikaidreieck. 
Unterseite der Schenkel stärker bedornt (> 8 Dornen), Vorderschenkel über der 
ganzen Länge mit einer Reihe von 12-15 kurzen Dornen (Enddornen nicht mit- 
gerechnet). Subgenitalplatte des 6 fast symmetrisch, mit zwei gleich grossen, 
fingerförmigen Styli.] 

3 Discus des Pronotums mit variabler Zeichnung aber ohne parallele 
dunkle Längsstreifen (Abb. 2); falls Längsstreifen angedeutet sind, 

divergieren diese nach hinten deutlich (Abb. 3) [Ectobius- Arten] 4 

Discus des Pronotums mit zwei ± parallelen, dunklen Längsstreifen 
(Abb. 1) Blattella germanica (Linnaeus) - Deutsche Schabe 

[Flügel reichen beim o* nicht ganz bis zur Spitze des Abdomens, beim 9 über- 
ragen sie es oft um 3-4 mm. Hinterflügel in Ruhelage ca. 0.5 mm kürzer als 
Deckflügel. Analfeld der Hinterflügel mit gefaltetem Apikaidreieck. Unterseite 
der Schenkel stärker bedornt (> 8 Dornen), Vorderschenkel über der ganzen 
Länge mit einer Reihe von 11-14 kurzen Dornen (Enddornen nicht mitge- 
rechnet). Subgenitalplatte des o* im Umriss ± rechteckig, mit zwei unter- 



ECTOBIUS VITTIVENTRIS (BLATTODEA) 



403 



mw 



i j 



sg 



/ 



1 

st X ( -: 



3 



i 



dg 



.'-#*o « 



>'*sf" ; •■^-•4^' ' 



'H'l*'' 



• 



-, 




i sg 




» 












8 



r— ^ 



..--' 

.» V^ 



; .. 




Abb. 6-8 
Abdomenende S, ventral: (6) Blattella germanica von Bern/BE. Abdomen <?. dorsal: (7) 
Ectobius vittiventris von Riehen/BS, (8) E. lucidus von Aesch/BL. Abkürzungen: c = Cercus, dg 
= Drüsengrube, s = Supraanalplatte, sg = Subgenitalplatte, st = Stylus. Massstab 1 mm. 
Zeichnungen A. Coray. 



404 H. BAUR ET AL. 

schiedlich grossen, seitlich gelegenen Sty li (Abb. 6); Subgenitalplatte wird bei 
beiden Geschlechtern von der Supraanalplatte völlig bedeckt und weit überragt 
(Abb. 6).] 

4 Discus des Pronotums ± einfarbig bernsteinfarben, gelegentlich mit 
unauffälligen, dunklen Flecken (Abb. 2, 15, 16). Deckflügel stellenweise 
fein gepunktet, aber ohne grössere, dunkle Flecken (Abb. 4); Flügel 
überragen das Abdomenende in beiden Geschlechtern um 0.3-3.1 mm 

Ectobius vittiventris (A. Costa) - Bernstein- Waldschabe 

[Grosse Ectobius-Art, Gesamtlänge o* 12.4-14.6 mm, 9 9.3-11.9 mm. 
Drüsengrube queroval, nur an den Rändern mit Haaren besetzt, in der Mitte kahl 

und höchstens mit schwacher Längserhebung, aber ohne Zäpfchen (Abb. 7, 10- 
12). Stylus des 6 relativ gross, oberseits mit einem Streifen dicht anliegender 
Haare (Abb. 7).] 

Discus des Pronotums mit auffälliger, dunkler Zeichnung (Abb. 3) oder 

ganz dunkel (Abb. 17, 19, 20), falls hell (Abb. 18), dann Deckflügel 

meist mit deutlichen dunklen Flecken (Abb. 5); Flügel lassen beim 9 

das Abdomenende unbedeckt 5 

5 Discus des Pronotums ohne betonte Hinterecken, ± gleichmässig ocker- 
farben bis dunkelbraun (Abb. 17, 18). Drüsengrube klein, queroval, in 
der Mitte mit einem dicht mit Haaren besetzten, zweigipfligen Zäpfchen 
(Abb. 13); Stylus des cT gross, ähnlich demjenigen von E. vittiventris in 

Abb. 7 Ectobius lapponicus (Linnaeus) - Lappland-Waldschabe 

Discus des Pronotums mit betonten Hinterecken, entweder in der Mitte 
aufgehellt (Abb. 3) oder gleichmässig braunschwarz bis schwarz (Abb. 
19, 20). Drüsengrube gross, nur leicht queroval bis fast halbkreisförmig, 
in der Mitte zerstreut mit Haaren besetzt, aber ohne Erhebung (Abb. 8); 
Stylus des 6 klein (Abb. 8) 6 

6 Discus des Pronotums in der Mitte ± aufgehellt (Abb. 3). Deckflügel 
beim 9 leicht verkürzt, aber Länge > 6 mm 

Ectobius lucidus (Hagenbach) - Hagenbachs Waldschabe 

[Grössere, stärker aufgehellte Art, Länge des Pronotums S 2.4 mm, 9 2.6 mm, 
der Deckflügel 6 10.7-11.3 mm, 9 6.3-6.5 mm (Durchschnittswerte ver- 
schiedener Fundorte aus der Schweiz und Deutschland, nach Bohn, 1989: 323). 
Drüsengrube leicht queroval (Abb. 8).] 

Discus des Pronotums einfarbig dunkelbraun bis schwarz, meist ohne 
Aufhellung in der Mitte (Abb. 19, 20); Deckflügel beim 9 stärker 

verkürzt, Länge < 6 mm Ectobius sylvestris (Poda) - Echte Waldschabe 

[Kleinere, sehr dunkle Art, Länge des Pronotums 8 2.0-2.2 mm, 9 2.3-2.4 mm, 
der Deckflügel S 9.7-11.6 mm, 9 4.8-5.5 mm (Durchschnittswerte verschie- 
dener Fundorte aus der Schweiz und Deutschland, nach Bohn 1989: 323). 
Drüsengrube fast halbrund, etwas grösser als bei E. lucidus (vgl. Bohn, 1989: 
327, Abb. 5). Achtung: gelegentlich haben die 9 9 ebenfalls ein aufgehelltes 
Pronotum und können deshalb leicht mit E. lucidus verwechselt werden.] 

7 Flügel verkürzt, beim â das Abdomenende freilassend, beim 9 schup- 
penartig, sich nicht berührend. Pronotum einheitlich schwarzbraun 

Blatta orientalis Linnaeus - Küchenschabe 

Flügel überragen das Abdomenende in beiden Geschlechtern deutlich. 
Pronotum braun, ± aufgehellt 8 



ECTOBIUS VirriVENTRIS (BLATTODEA) 



405 




Abb. 9 
Oothek: Ectobius vittiventris von Bottmingen/BL. Massstab 1 mm. Zeichnung A. Coray. 

Deckflügel einfarbig braun. Dunkle Zeichnung des Pronotums häufig 
verwaschen, ± gelblichweiss umrandet 

Periplaneta americana (Linnaeus) - Amerikanische Schabe 

Vorderrand der Deckflügel mit elfenbeinfarbenem Streifen, ansonsten 
braun. Dunkle Zeichnung des Pronotums kontrastreicher und schärfer 
gegen die gelblichweisse Umrandung begrenzt 
Periplaneta australasiae (Fabricius) - Australische Schabe 



Verbreitung 

In der Schweiz ist E. vittiventris sowohl auf der Nord- als auch auf der Südseite 
der Alpen verbreitet (Abb. 21). Im Norden kommt die Art im zentralen und östlichen 
Mittelland und im Genferseegebiet vor, wobei viele Verbreitungslücken bestehen. Es 
ist zu beachten, dass die Häufung von Daten in der Umgebung von Bern, Basel, Luzern 
und besonders Zürich (vgl. Appendizes) allein auf die besonderen Aktivitäten der 
Autoren und des Ungezieferbestimmungsdienstes des NML zurückzuführen ist! 
Bemerkenswert ist das fast vollständige Fehlen von älteren Funddaten auf der 
Alpennordseite. Mit der Ausnahme von einem Exemplar von Küsnacht (ZH) aus dem 
Jahr 1961 sowie einem undatierten, aber wahrscheinlich in den 50er Jahren gesam- 
melten Männchen von Basel (BS), stammen alle Angaben aus der Zeit nach 1985. Mit 
Hilfe einer linearen Regression konnte ferner festgestellt werden, dass der Anteil an E. 
vittiventris-Meldungen bei den UGZ-Daten signifikant zunahm (b = 0.704, r 2 = 0.518, 
F = 8.598, p < 0.02, vgl. Abb. 22). In den letzten Jahren war E. vittiventris schliesslich 
die am häufigsten verzeichnete Tierart. 

Südlich der Alpen, besonders im Sottoceneri, ist E. vittiventris weit verbreitet. 
Viele Funde gehen hier auf ältere Angaben zurück. Die Feststellung von Fruhstorfer 
(1921, sub E. neolividus), E. vittiventris sei "eines der Charaktertiere des südlichen 
Tessin", hat durchaus noch seine Gültigkeit (Baur, pers. Beob.). 

Die Verbreitungskarte (Abb. 21) weist einige markante Lücken auf. So scheint 
die Art im Jura und auf der Alpennordseite im Wallis und Graubünden weitgehend zu 



406 



H. BAURET AL. 





12 



13 




14 



Abb. 10-14 
Drüsengrube: (10) Ectobius vittiventris von Chavannes-près-Renens/VD, (11) Lancy/VD, (12) 
Winterthur/ZH, (13) E. lapponicus von Eclépens/VD. Linker Phallomer, apikaler Teil: (14) E. 
vittiventris von Chavannes-près-Renens/VD. Massstab 0.5 mm. Fotos H. Baur. 



fehlen. Nur zwei Exemplare mit der unspezifischen Angabe "Wallis" befinden sich in 
CAN bzw. MHNG. Aus dem Oberengadin (GR) lag uns ein einzelnes Männchen von 
Pontresina aus der Sammlung CAN vor. Dem Exemplar fehlt allerdings das Abdomen, 
ansonsten stimmt es mit der Diagnose der Art aber einwandfrei überein. 

E. vittiventris bevorzugt in der Schweiz die kolline und submontane Höhenstufe 
bis ca. 600 m ü.M. (Abb. 23). Nur an wenigen Orten steigt die Art nördlich der Alpen 
auf ca. 1000 m ü.M. (Engelberg, OW). Im Süden ist sie gelegentlich in noch höheren 



ECTOBIUS V1TTIVENTRIS (BLATTODEA) 



407 




Abb. 15-20 
Imagines im natürlichen Lebensraum: (15) Ectobius vittiventris <S und (16) 9 von Riehen/BS. 
(17) E. lapponicus S und (18) 9 von Wehr/Baden-Württemberg, (19) E. sylvestris 3 von 
Beurnevésin/JU und (20) 9 von Wehr/Baden-Württemberg. Fotos A. Coray. 

Lagen anzutreffen, z. B. wurde sie am Passo Pairolo (TI) auf 1406 m ü.M. von Fruh- 
storfer gesammelt. Der Einzelfund auf 1800 m ü.M. betrifft das zuvor erwähnte 
Männchen von Pontresina. 

Mehrere Funde von E. vittiventris wurden uns aus Baden-Württemberg, 
Deutschland, mitgeteilt (Appendix 2). Die Art kommt dort entlang des Rheins an 
mehreren Stellen bis Gottenheim bei Freiburg vor 1 . 



1 Das Manuskript war bereits im Druck, als uns 2 Exemplare aus einer Wohnung in 
Stuttgart, ca. 130 km nordöstlich von Gottenheim, vorgelegt wurden (leg. Oktober 2003, 1 9 
coll. NMBE, 1 5 coli. Matthias Schöller, Berlin). 



408 



H. BAURET AL. 




Abb. 21 
Verbreitung von Ectobius vittiventris in der Schweiz. O = Funde vor 1986 (n = 48), # = Funde 
nach 1985 (n = 536), ♦ = Funde nur durch UGZ-Daten (n = 1637) belegt (vgl. Material und 
Methoden). 

Phänologie 

E. vittiventris kann das ganze Jahr über angetroffen werden. Einen Überblick 
zum jahreszeitlichen Erscheinen gibt die Abb. 24. Die MUS- und UGZ-Daten zeigen 
eine recht gute Übereinstimmung. In beiden Fällen wurde die grösste Individuendichte 
nach einem steilen Anstieg im August registriert; doch bereits im September gingen die 
Zahlen stark zurück und nahmen bis Dezember kontinuierlich ab. In den Monaten 
Januar bis April wurden nur vereinzelt Individuen gesammelt. Eine Aufteilung der 
MUS-Daten nach Geschlecht bzw. Stadium (Abb. 25) zeigte, dass in diesen Monaten 
ausschliesslich ältere Nymphen (etwa 3.-5. Stadium) gefunden wurden. Die ersten 
Imagines traten in geringer Anzahl erst ab Ende Mai auf. In den folgenden Monaten 
nahmen zuerst die Männchen rasch zu und erreichten ihren Höhepunkt im Juli. Das 
Maximum der Weibchen befand sich rund einen Monat später im August, ab 
September nahmen aber auch sie schnell wieder ab. Parallel dazu stieg der Anteil 
gefundener Weibchen mit Oothek von Ende Juni bis in den Oktober. Vereinzelt 
konnten Imagines schliesslich noch bis in den November und Dezember hinein 
beobachtet werden. 

Die Verteilung der einzelnen Nymphenstadien wurde nicht im Detail unter- 
sucht. Wir haben jedoch festgestellt, dass im Winter meist mittlere und grössere 
Nymphen (ca. 4-7 mm) gefunden wurden. Im Juli konnten gelegentlich noch recht 
kleine Nymphen gesammelt werden (Bern, Obstberg, 23.7.2000, leg. H. Baur). Einige 
dieser Exemplare waren deutlich kleiner (2-3 mm, ev. 2. Stadium) als jene vom 
Winterhalbjahr. 



ECTOBIUS VimVENTRIS (BLATTODEA) 



409 




1991 



1992 1993 



1994 



1995 1996 
Jahr 



1997 1998 1999 



2000 



Abb. 22 
Lineare Regression des Anteils (in %) von Ectobius vittiventris an der Gesamtheit aller durch 
den UGZ erfassten Tierarten. Die Steigung der Regressionsgerade für die Jahre 1991-2000 ist 
signifikant (b = 0.704, p < 0.02; n Ev = 1640, n = 21593). 



Verhalten 

Hierzu können nur Einzelbeobachtungen mitgeteilt werden. E. vittiventris ist 
tagsüber am ehesten in seinen Verstecken anzutreffen, z. B. unter Blumentöpfen, in 
Rollladenkästen, im Laub oder unter Steinen. Werden die Tiere gestört, suchen sie so- 
fort die nächste Spalte auf. Die Männchen benutzen auf der Flucht oft ihre Flügel, 
wobei sie im Flug auch an Höhe gewinnen können. Demgegenüber sind die Weibchen 
höchstens zu einem Sinkflug fähig. In Gebäuden wird E. vittiventris oft auch am Tag 
an ungeschützten Orten angetroffen, während die Art im Freien normalerweise erst in 
der Dämmerung aktiv wird. Die Tiere krabbeln dann aus ihren Verstecken hervor und 
klettern auf niederen Zweigen und Laubwerk im Gestrüpp von Waldrändern oder 
städtischen Grünanlagen umher. Mehrmals konnte beobachtet werden, wie sich 
mehrere Männchen gegenseitig vertrieben, besonders in Anwesenheit eines Weib- 
chens. Vor der Paarung nähert sich das Männchen dem Weibchen rückwärts mit fast 
vertikal erhobenen, aber zusammengefalteten Flügeln und bietet so dem Weibchen 
seine Drüsengrube dar. Bei der endgültigen Kopulationsstellung weisen dann die 
Köpfe des Pärchens in entgegengesetzte Richtungen. Im Spätsommer und Herbst 
wurden gelegentlich Weibchen mit Ootheken angetroffen (s. oben). Wo und unter 
welchen Umständen die Ootheken abgelegt werden, Hess sich nicht feststellen. 
Nymphen unterschiedlicher Grösse wurden regelmässig zusammen mit Imagines auf 
Blättern und Zweigen, aber auch in Häusern angetroffen. 



410 



H. BAURET AL. 



300 



250 



c 200 

3 

"2 
> 

C 



150 



c 
< 



100 



50 



Alpennordseite 
■ Alpensüdseite 



■ I 



// 



300 400 500 600 700 800 900 1000 1100 1200 1300 1400 
Höhe über Meer in rn 



Abb. 23 
Höhenverbreitung von Ectobius vittiventris in der Schweiz aufgrund der MUS-Daten 
(Alpennordseite n = 516, Alpensüdseite n = 67). Der Einzelfund auf 1800 m über Meer wurde 
weggelassen. 



Fundumstände 

E. vittiventris konnte im Freien besonders in Hecken und entlang von Wald- 
rändern auf Sträuchern beobachtet werden. Ein Blick auf die MUS-Daten (Appendix 
2) zeigt jedoch, dass in der Nordschweiz die allermeisten Funde (> 85%) in oder um 
Häuser gemacht wurden. Bei den UGZ-Daten dürfte dieser Anteil nahezu 100% 
betragen. Dies ist nicht weiter verwunderlich, kamen die Mitarbeitenden des UGZ - 
entsprechend ihrer Tätigkeit - doch nur an Material von solchen Orten heran! Die 
Daten sind daher ganz offensichtlich mit einem systematischen Fehler behaftet. Da von 
uns keine gezielten Untersuchungen zum Auftreten dieser Art ausserhalb des 
Siedlungsbereichs durchgeführt wurden, wird hier auf eine statistische Analyse der 
Fundumstände verzichtet. 

Obwohl E. vittiventris oft in Häuser eindringt, haben wir keine Hinweise auf 
eine mögliche Vermehrung in Gebäuden erhalten. Selbst bei grösseren Vorkommen, 
wie in Bottmingen, Bruderholz (BL, vgl. Appendix 2), handelte es sich lediglich um 
Exemplare, die über einen längeren Zeitraum von aussen eindrangen. Nach unseren 
Beobachtungen kann E. vittiventris nicht länger als ein paar Tage in geschlossenen 
Räumen überleben. 



ECTOBIUS VITTIVENTR1S (BLATTODEA) 



41 



40 



30 

c 
c 

Q> 

-a 

> 20 



-C 
■X 
N 

«I 



^C 






I 



■ UGZ 
MUS 




Jan Feb Mär Apr 



Mai Jun Jul Aug Sep Okt Nov Dez 
Monat 



Abb. 24 
Phänologie von Ectobius vittiventris in der Schweiz nach UGZ- (n = 1637) bzw. MUS-Daten 
(n = 569). Zum besseren Vergleich erfolgte eine Standardisierung der Anzahl Individuen in %. 



DISKUSSION 

Eine Analyse der Verbreitungsdaten (vgl. Abb. 21) macht deutlich, dass Ecto- 
bius vittiventris auf der Alpennordseite fast ausschliesslich nach Mitte der Achtziger- 
jahre und besonders in städtischen Gebieten gefunden wurde. Dadurch könnte der 
Eindruck entstehen, dass die Art wahrscheinlich aus dem Süden eingeschleppt wurde 
und sich danach rasant in den urbanen Regionen ausbreitete. In eine ähnliche Richtung 
weist die Zunahme des Anteils von E. vittiventris an der Gesamtheit aller UGZ-Daten 
von 1991-2000 (vgl. Tab. 1, Abb. 22). Obwohl eine Verschleppung von Ectobius-Arten 
schon mehrfach dokumentiert wurde (Hoebecke & Nickle, 1981; Chandler, 1985, 
1992), kann dieses Szenario durch unsere Daten jedoch nicht eindeutig belegt werden. 
Das Hauptproblem besteht darin, dass keine älteren Daten zum Vergleich heran- 
gezogen werden können. Fruhstorfer (1921) konzentrierte sich in der bisher einzigen 
faunistischen Bearbeitung der einheimischen Ectobius- Arten besonders auf das Tessin, 
das Wallis und die Bündner Südtäler. Seine Fundortsangaben für die Nordschweiz 
beruhten zumeist auf unüberprüfbaren Meldungen älterer Autoren. Zudem hatte er 
viele Arten nachweislich falsch bestimmt (Ramme, 1923, 1951; Zeuner, 1931; Harz, 
1957b). Die geringe Beachtung durch Entomologen führte ferner dazu, dass 
Waldschaben in Sammlungen generell unterrepräsentiert sind. Aufgrund des Fehlens 



412 H. BAUR ETAL. 

älterer Meldungen oder Belege kann somit nicht zwingend auf eine rezente Areal- 
erweiterung von E. vittiventris geschlossen werden. Es könnte durchaus sein, dass die 
Art früher sehr wohl vorhanden war, aber verkannt wurde. Hierfür sprechen die beiden 
Exemplare von Basel (BS) und Küsnacht (ZH) aus den Fünfziger- bzw. Sechziger- 
jahren (Appendix 2). Demgegenüber lässt insbesondere die Regressionsanalyse der 
UGZ-Daten (vgl. Abb. 22) den Schluss zu, dass die Grösse der Populationen von E. vit- 
tiventris in den letzten Jahren stark zugenommen hat. Allerdings könnte dagegen 
eingewendet werden, dass der Anstieg der Meldungen z. B. auf die zunehmende 
Beachtung in den Medien zurückzuführen sei. Dieser Effekt hat wahrscheinlich eine 
gewisse Rolle gespielt, wobei der UGZ nicht zuletzt durch die eigenen Aktivitäten - 
eine öffentliche Veranstaltung zum Thema Waldschaben im Oktober 1999, Merkblätter 
im Internet, etc. - mitgeholfen hatte, das öffentliche Interesse an der Schabe zu wecken. 
Das Ergebnis der Regression allein diesem Umstand zuzuschreiben, scheint uns jedoch 
nicht gerechtfertigt. Denn dort, wo E. vittiventris in den Wohnbereich eindringt, wird 
dieser Art wegen ihrer Ähnlichkeit mit der Deutschen Schabe Blattella germanica 
schnell einmal Beachtung geschenkt. Deshalb wäre ein Auftreten im gegenwärtigen 
Ausmass - E. vittiventris ist seit 1998 die am häufigsten registrierte Tierart am UGZ - 
auch in früheren Jahren bemerkt worden und hätte zumindest zu einer weitaus 
grösseren Zahl von Belegen in Sammlungen führen müssen. 

Bei mitteleuropäischen Insekten (Klausnitzer, 1982) und Spinnentieren (Thaler 
& Knoflach, 1995) sind Zunahmen von Populationen oder gar Arealerweiterungen ver- 
schiedentlich dokumentiert worden. Als Beispiel sei die Südliche Eichenschrecke 
Meconema meridionale (A. Costa) (Ensifera: Meconematidae) herausgegriffen, weil 
hier einige Parallelen zu E. vittiventris zu finden sind. Bei den Heuschrecken ist die 
faunistische Bearbeitung aber vergleichsweise gut (Thorens & Nadig, 1997), weshalb 
hier weitergehende Schlüsse möglich waren. Noch bis Mitte der Achtzigerjahre war die 
vorwiegend mediterrane M. meridionale nur von wenigen Orten nördlich der Alpen 
bekannt, in der Schweiz aus dem Tessin, Genferseebecken und unteren Wallis (Nadig, 
1981). Im Laufe von wenigen Jahren tauchte die Art dann plötzlich in den meisten 
Städten in der Nordschweiz auf (Thorens & Nadig, 1997). In Bern, Winterthur und 
Zürich konnte sie ab Mitte der Neunzigerjahre sogar zusammen mit E. vittiventris 
beobachtet werden (Baur, unpubl.). Da M. meridionale in der älteren Literatur für diese 
Orte nicht erwähnt wird, erscheint eine jüngere Arealerweiterung wahrscheinlich 
(Thorens & Nadig, 1997). Hinzu kommt, dass die Art und Weise der Ausbreitung durch 
passive Verschleppung teilweise bekannt ist (Tröger, 1986). Nach Klausnitzer (1982) 
können sich wärmeliebende Arten wie M. meridionale in Städten besonders gut halten, 
weil diese "Wärmeinseln" durchschnittlich höhere Temperaturen aufweisen als die 
umliegenden, ländlichen Regionen. Falls E. vittiventris sich wirklich rezent nach 
Norden ausgebreitet hätte, wäre auch diese Art am ehesten in Städten zu erwarten. 
Unsere Daten lassen freilich auch in dieser Hinsicht keinen eindeutigen Schluss zu, da 
wir nicht wissen, inwieweit das Vorkommen von E. vittiventris tatsächlich auf urbane 
Regionen beschränkt ist. 

Hinsichtlich der Gesamtverbreitung bleibt abzuklären, wie weit E. vittiventris 
ins Walliser Rhonetal vordringen konnte. Wie bereits erwähnt, sind nur zwei Exem- 
plare mit der unspezifischen Bezeichnung "Wallis" bekannt (vgl. Appendix 2). Diese 



ECTOBIUS VirnVENTRJS (BLATTODEA) 



413 



100 



80 - 



c 

3 60 



40 



20 



/ 



Nymphe 

Männchen 

A -Weibchen 

•' \ Oothek 




Jan Feb Mär Apr Mai Jun Jul Aug Sep Okt Nov Dez 

Monat 

Abb. 25 
Phänologie von Ectobius vittiventris nach Geschlecht bzw. Stadium (MUS-Daten, n = 596). 



könnten z. B. auch von der Alpensüdseite oder dem untersten Wallis an der Grenze 
zum Kanton Waadt stammen. Demgegenüber ist das Auftauchen von E. vittiventris in 
Baden- Württemberg (vgl. Appendix 2) angesichts der grenznahen Basler Funde nicht 
überraschend. Die Art wurde für Deutschland aber noch nicht gemeldet (Harz, 1957b; 
Bonn, 2000). 

Die überwiegend kolline bis montane Höhenverbreitung von E. vittiventris 
entspricht derjenigen der meisten anderen Ectobius- Arten, einzig E. sylvestris ist 
regelmässig in subalpinen und alpinen Lagen anzutreffen (Harz, 1957b; Baur, Coray, 
unpubl). Der Fund eines einzelnen Männchens von E. vittiventris in Pontresina auf 
1800 m ü.M. fällt somit völlig aus dem Rahmen. Hier wäre eine unbeabsichtigte 
Verschleppung aus dem nahen Bergeil, wo die Art häufig ist, denkbar. 

Zur Anzahl der Nymphenstadien und zum Entwicklungszyklus konnten wir 
keine direkten Beobachtungen machen. Bei einigen Ectobius-Aiiten wurden fünf oder 
sechs Nymphenstadien festgestellt (Brown, 1973c, 1980), was auch für E. vittiventris 
zutreffen dürfte. Die Phänologie der Imagines, Nymphen und Ootheken (Abb. 25) 
macht ferner eine zweijährige Entwicklung wahrscheinlich. Das Auffinden von älteren 
Nymphen im Winter ist ein klarer Hinweis dafür, dass die Postembryonalentwicklung 
nicht in einem Jahr abgeschlossen werden kann. Der Lebenszyklus könnte daher etwa 
folgendermassen aussehen: Nach Ablage der Oothek im Sommer und Herbst erfolgt 
der Schlupf der Nymphen im darauf folgenden Frühjahr. Ein Indiz hierfür sind kleine 



414 H. BAURET AL. 

Nymphen (höchstens 2. Stadium), welche fast ausschliesslich im Juli gefunden 
wurden. Der genaue Zeitpunkt des Schlupfes konnte jedoch nicht bestimmt werden, 
könnte aber ungefähr im Mai oder Juni liegen. Die Nymphen wachsen anschliessend 
bis zum Herbst langsam heran und überwintern in etwas älteren Stadien. Unsere Funde 
von grösseren Nymphen in den Wintermonaten machen dies deutlich. Das Alter dieser 
Nymphen konnten wir nur ungefähr abschätzen, sie befanden sich jedoch mindestens 
im 2. oder 3. Stadium. Nach dem zweiten Winter reifen die Nymphen schliesslich von 
Ende Mai bis August zur Imago. Die Imaginalhäutung findet bei den Männchen 
wahrscheinlich überwiegend im Juli statt, da in diesem Monat die grösste Anzahl von 
Imagines gefunden wurde (vgl. Abb. 25). Die Weibchen werden durchschnittlich etwa 
einen Monat später adult. 

Für E. lapponicus, E. pallidas und E. sylvestris wurde bereits ein zweijähriger 
Entwicklungszyklus festgestellt (Morvan, 1972; Brown, 1973c, 1980). Brown (1973c, 
1980) zeigte, dass eine Diapause bei niederen Temperaturen für die Eientwicklung von 
E. pallidus und E. lapponicus obligatorisch ist. Die Überwinterung der Nymphen 
erfolgte bei E. pallidus im 2. bis 5. Stadium (Brown, 1980). Bei E. lapponicus konnte 
Brown (1973c) eine Quieszenz im 2. oder 3. sowie eine fakultative Diapause im 4. 
Nymphenstadium nachweisen. Erfolgte die Aufzucht bei einer konstanten Temperatur 
von 20 °C, so konnten sich die Nymphen nicht bis zur Imago entwickeln. In England 
(Brown, 1973c), Frankreich (Morvan, 1972) und Tschechien (Holusa & Kocârek, 
2000) vollzogen die meisten Individuen die Imaginalhäutung bereits im Mai und Juni. 
E. lapponicus ist wie E. pallidus (vgl. Morvan, 1972; Brown, 1973c, 1980) also rund 
ein bis zwei Monate früher adult als E. vittiventris. Der Entwicklungszyklus stimmt bei 
dieser Art somit besser mit demjenigen von E. sylvestris in Frankreich überein 
(Morvan, 1972: 271, Fig. 12). 

Zweifellos haben diese Angaben zur Entwicklung von E. vittiventris einen 
etwas vorläufigen Charakter. Sie beruhen aber auf der bestmöglichen Interpretation 
unserer Daten. Experimentelle Untersuchungen, wie sie von Brown (1973c, 1980) in 
exemplarischer Weise bei E. lapponicus und E. pallidus durchgeführt wurden, wären 
zur Überprüfung der aufgestellten Hypothese sicher zu begrüssen. 

Unsere Beobachtungen zum Verhalten von E. vittiventris entsprechen ungefähr 
dem gattungstypischen Muster. Besonders das Paarungsverhalten wird in ähnlicher 
Weise auch von E. lapponicus geschildert (Ramme, 1920; Harz, 1960). Im Unterschied 
zu den überwiegend tagaktiven E. lapponicus und E. sylvestris (vgl. Harz, 1957b, 
1960; Dreisig, 1971) scheint E. vittiventris im Freien vor allem in der Dämmerung und 
nachts aktiv zu sein. In Häusern ist die Art aber auch tagsüber überall anzutreffen. 
Damit unterscheidet sie sich in ihrem Verhalten auch von Blattella germanica, welche 
nur nachts aus ihren Verstecken hervorkriecht und sich besonders an feucht-warmen 
Orten (z. B. beheizte Badezimmer, Küchen) aufhält. Ferner sind beide Geschlechter 
dieser Art flugunfähig, während die Männchen von E. vittiventris über kürzere 
Strecken fliegen können. 

Auch unter Berücksichtigung eines systematischen Fehlers wird deutlich, dass 
E. vittiventris relativ häufig in und um Häuser angetroffen wird (vgl. Appendix 2). Dies 
führte teilweise zu Verwirrung, da diese Art zumeist mit Blattella germanica verwech- 
selt wurde. Wie wir bereits früher festgestellt haben (Landau Lüscher et ah, 2003), 



ECTOBIUS VITTWENTRIS (BLATTODEA) 415 

muss E. vittiventris jedoch nicht bekämpft werden, da die Tiere in Gebäuden offen- 
sichtlich nicht mehr als ein paar Tage überleben können. Möglicherweise verhindert 
auch eine obligatorische Diapause in der Entwicklung der Eier bzw. der Nymphen, wie 
sie bei anderen Ectobius-Aiten festgestellt wurde (Brown, 1973c, 1980), eine Fort- 
pflanzung bei Raumtemperatur. Eine Bekämpfung ist daher nicht nur nicht angezeigt, 
sondern auch nutzlos, da immer wieder neue Exemplare von aussen eindringen. 

Das Auftreten von Waldschaben in Gebäuden ist seit längerem bekannt (Weid- 
ner, 1972; Abraham, 1979; Jonge, 1984; Chandler, 1985). Ferner beschreibt Mielke 
(2000) in einer neueren Arbeit einen Massenbefall von E. lapponicus in einem Spital 
in Magdeburg (Deutschland). Allerdings handelte es sich bei all diesen Ereignissen 
jeweils um Einzelfälle. Im Gegensatz dazu tritt E. vittiventris nun schon seit über zehn 
Jahren regelmässig in Siedlungen in der Schweiz auf. Es ist somit wahrscheinlich, dass 
die Art sich an diesen Orten bereits etabliert hat. 

DANKSAGUNG 

An erster Stelle gebührt unser Dank Horst Bohn (Zoologisches Institut der 
Universität, München, Deutschland), der uns während Jahren auf vielfältige Weise 
unterstützte und die mikroskopischen Präparate in Abb. 10-14 anfertigte. Elsa Obrecht 
(NMBE), Alain Jacob (Bern, Schweiz) und ein anonymer Begutachter lasen das 
Manuskript kritisch durch und gaben uns wertvolle Anregungen. Adolf Nadig (Chur, 
Schweiz) ermöglichte dem Erstautor die Durchsicht seiner Sammlung (CAN), noch 
bevor diese im Oktober 2001 dem MHNG übergeben wurde. Für den freundlichen 
Empfang in Chur und die Gastfreundschaft sei ihm und seiner Frau herzlich gedankt. 
Für die Ausleihe von Material oder Angaben zu Exemplaren in Sammlungen danken 
wir den folgenden Personen und Institutionen: Georg Artmann (CGA), Wolf gang 
Billen (Pflanzengesundheitsdienst, Amt für Landwirtschaft, Lörrach, Deutschland), H. 
Bohn (München), Daniel Burckhardt (NHMB), Peter Herger (NML), Charles Lienhard 
(MHNG), Andreas Müller (ETHZ), Isolde Dorandt und Michael Ohi (ZMB), sowie 
sämtlichen Sammlerinnen und Sammlern. 

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Zoologischen Museums Hamburg 6: 229-230. 
Beier, M. 1967. Schaben. Die neue Brehm-Bücherei 379, Ziemsen, Wittenberg Lutherstadt, 

38 pp. 
Besuchet, C. 1956. Biologie, morphologie et systématique des Rhipidius. Mitteilungen der 

schweizerischen entomologischen Gesellschaft 29: 73-144. 
Bohn, H. 1989. Revision of the sylvestris group of Ectobius Stephens in Europe (Blattaria: 

Blattellidae). Entomologica scandinavica 20: 317-342. 
Bohn, H. 2000. Blattoptera - Schaben (pp. 105-109). In: Hannemann, H.-J., Klausnitzer, B. & 

Senglaub, K. (Hrsg.). Exkursionsfauna von Deutschland, Band 2. Wirbellose: Insekten 

(9. Auflage). Spektrum, Heidelberg, 959 pp. 
Brown, V. K. 1973a. The biology and development of Brachy gaster minutus Olivier 

(Hymenoptera: Evaniidae), a parasite of oothecae of Ectobius spp. (Dictyoptera: 

Blattidae). Journal of Natural History 7: 665-674. 



416 H. BAURET AL. 



Brown, V. K. 1973b. Aspects of reproductive biology of three species of Ectobius (Dicty optera: 

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(Dictyoptera: Blattellidae). Entomological News 96: 98-100. 
Chandler, D. S. 1992. New records of Ectobius lapponicus in North America (Dictyoptera: 

Blattellidae). Entomological News 103: 139-141. 
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Harz, K. 1957a. Zur Biologie der Waldschabe Ectobius silvestris (Poda) (Blattopteroida, 
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Harz, K. 1957b. Die Geradflügler Mitteleuropas. Gustav Fischer, Jena, XXIII + 494 pp., 10 pi. 

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Jonge, J. T. de 1984. Opmerkelijke insekten in en om gebouwen in 1983. Entomologische 
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Klausnitzer, B. 1982. Grossstädte als Lebensraum für das mediterrane Faunenelement. Ento- 
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ECTOBIUS VITTIVENTRIS (BLATTODEA) 417 



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29-36. 



418 



H. BAUR ET AL. 



Appendix 1: UGZ-Daten (vgl. Material und Methoden) der Jahre 1991-1999 und 2001 von 
Ectobius vittiventris. Aufgelistet ist die Anzahl Datensätze (N) pro Ort, sortiert nach Kanton (Kt) 
und Ort. Die fragliche Meldung von Inden (VS) konnte nicht mehr überprüft werden und wurde 
in der Auswertung (Tab. 1, Abb. 21, 22, 24) weggelassen. 



N Ort 



Kt 



N Ort 



Kt 



N Ort 



Kt 



2 Aarau 


AG 


4 Birmensdorf 


ZH 


1 Bremgarten 


AG 


2 Bonstetten 


ZH 


1 Ennetbaden 


AG 


1 Brütten 


ZH 


1 Jonen 


AG 


1 Brüttisellen 


ZH 


1 Mellingen 


AG 


3 Buchs 


ZH 


1 Nussbaumen b. Baden 


AG 


5 Bülach 


ZH 


2 Rudolfstetten 


AG 


1 Dielsdorf 


ZH 


1 Spreitenbach 


AG 


3 Dietikon 


ZH 


1 Sterten 


AG 


3 Dietlikon 


ZH 


1 Suhr 


AG 


2 Dinhard 


ZH 


1 Unterehrendingen 


AG 


1 1 Dübendorf 


ZH 


1 Wettingen 


AG 


1 Dürstelen 


ZH 


1 Widen 


AG 


1 Effretikon 


ZH 


1 Würenlos 


AG 


1 Egg b. Zürich 


ZH 


1 Zufikon 


AG 


1 Eglisau 


ZH 


1 Bern 


BE 


1 Elgg 


ZH 


1 Ipsach 


BE 


6 Erlenbach 


ZH 


1 Kilchberg 


BL 


2 Esslingen 


ZH 


1 Oberwil 


BL 


1 Fällanden 


ZH 


1 Reinach 


BL 


2 Fällanden, Benglen 


ZH 



3 Collonge-Bellerive GE 
Lancy, Grand-Lancy GE 
Inwil LU 

2 Meggen LU 

Stans NW 

Altstätten SG 

Amden SG 

2 Jona SG 

4 Schaffhausen SH 
Dornach SO 
Feldbrunnen SO 
Altendorf SZ 
Freienbach, Pfäffikon SZ 
Schwyz SZ 
Schwyz, Ibach SZ 
Siebnen bei Wangen SZ 
Wollerau SZ 
Inden VS 
Baar ZG 
Cham ZG 

3 Zug ZG 
Adliswil ZH 
Aeugst am Albis ZH 

6 Affoltern am Albis ZH 

3 Bassersdorf ZH 

1 Bertschikon bei Uster ZH 



3 Fällanden, Pfaffhausen ZH 

1 Fehraltorf ZH 

2 Gattikon ZH 
7 Geroldswil ZH 

3 Glattbrugg ZH 
3 Gossau ZH 
2 Greifensee ZH 
2 Grüningen ZH 

1 Griit bei Wetzikon ZH 

2 Hausen am Albis ZH 

3 Herrliberg ZH 
2 Hinwil ZH 

2 Hombrechtikon ZH 

3 Horgen ZH 

1 Humlikon ZH 

2 Illnau ZH 
11 Kilchberg ZH 

7 Kloten ZH 

4 Küsnacht ZH 

5 Langnau am Albis ZH 
2 Männedorf ZH 

1 Maur, Binz ZH 
4 Maur, Ebmatingen ZH 

2 Maur, Forch ZH 

6 Meilen ZH 
9 Meilen, Feldmeilen ZH 



2 Mönchaltorf ZH 

4 Nürensdorf ZH 

4 Oberengstringen ZH 

1 Oberhasli ZH 
4 Oberrieden ZH 

2 Obfelden ZH 
1 Oetwil am See ZH 

1 Oetwil an der Limmat ZH 
4 Opfikon ZH 

2 Otelfmgen ZH 

3 Ottenbach ZH 
3 Regensdorf ZH 

3 Regensdorf, Watt ZH 
1 Richterswil ZH 

4 Rümlang ZH 

5 Rüschlikon ZH 
1 Russikon ZH 
1 Rüti ZH 

6 Schlieren ZH 
1 Schwerzenbach ZH 
1 Seegräben, Aathal ZH 

1 Stadel ZH 

2 Stäfa ZH 

3 Stäfa, Uerikon ZH 

1 Steinmaur ZH 

8 Thalwil ZH 

4 Uetikon am See ZH 
16 Uitikon- Waldegg ZH 

2 Unterengstringen ZH 
12 Urdorf ZH 

9 Uster ZH 

2 Volketswil ZH 
9 Wädenswil ZH 
1 Wädenswil, Au ZH 

7 Wallisellen ZH 

1 Weisslingen ZH 

3 Wernetshausen ZH 
7 Wettswil am Albis ZH 

2 Wetzikon ZH 
1 Winkel bei Bülach ZH 
1 Winterberg ZH 

18 Winterthur ZH 

11 Zollikon ZH 

10 Zollikon, Zollikerberg ZH 

7 Zumikon ZH 

1313 Zürich ZH 



ECTOBIUS VITTIVENTRIS (BLATTODEA) 



419 



Appendix 2: Vollständige Auflistung aller MUS-Daten (vgl. Material und Methoden) von 
Ectobius vittiventris, sortiert nach Land, Kanton, Ort und Datum. Bei Exemplaren bzw. 
Ootheken, die vermessen wurden, ist die Anzahl fett hervorgehoben. *Die Anzahl der Exemplare 
von Bottmingen, Bruderholz (BL) wurde aufgrund fehlender Genauigkeit beim Datum in der 
Auswertung (Resultate, Abb. 23-25) weggelassen. BW = Baden-Württemberg; CH = Schweiz; 
Coli. = Sammlung (Abkürzungen, siehe Material und Methoden); D = Deutschland; I = Italien; 
Kt = Kanton bzw. Region (Abkürzungen der Kantone der Schweiz, siehe Material und 
Methoden); Ld = Land; Mon = Monat; NI = Norditalien; Ny = Nymphe; Oo = Oothek; TO = 
Toskana. 



(J 


9 


\> 


Oo Ld 


Kt 


Ort 


Tag 


Mon 


Jahr 


Höhe Coli. 


Bemerkungen 


1 


1 





CH 


AG 


Aarau 


28 


6 


2000 


350 UGZ 


Altersheim, Zimmer 





1 





CH 


AG 


Aarau 


5 


10 


2000 


350 UGZ 







1 





CH 


AG 


Baden 


4 


8 


2000 


380 UGZ 










2 


CH 


AG 


Bergdietikon, Kindhausen 


29 


5 


2000 


581 UGZ 


aussen, unter Blumentöpfen 


1 








CH 


AG 


Jonen 


8 


6 


2000 


401 UGZ 


Haus 








3 


CH 


AG 


Umiken 


16 


3 


2000 


360 UGZ 


Haus 


2 





1 


CH 


AG 


Wettingen 


4 


8 


2000 


390 UGZ 


Haus 





2 





1 CH 


AG 


Wettingen 


20 


10 


2001 


390 NMBE 


Haus 





2 


1 


CH 


AG 


Zurzach 








339 UGZ 







2 





2 CH 


BE 


Bem 


30 


6 


2000 


550 UGZ 


Kinderkrippe, Sandkasten 





1 





CH 


BE 


Bem 


7 


11 


2001 


550 NMBE 


Haus 








1 


CH 


BE 


Bern 


16 


2 


2001 


550 NMBE 


Wohnung 





4 





CH 


BE 


Bern, Breitenrain 


10 


9 


1996 


550 NMBE 







,1 





CH 


BE 


Bern, Kirchenfeld 


21 


9 


2000 


560 NMBE 


Haus 


2 








CH 


BE 


Bern, Obstberg 


21 


6 


2000 


565 NMBE 


auf Sträuchem, nachts 





1 





CH 


BE 


Bern, Obstberg 


3 


7 


2000 


565 NMBE 


Umgebung Haus 








20 


CH 


BE 


Bem, Obstberg 


23 


7 


2000 


565 NMBE 


Hecke 





1 





CH 


BE 


Bern, Obstberg 


4 


8 


2000 


565 NMBE 


Wohnung 





1 





CH 


BE 


Bem, Obstberg 


23 


8 


2000 


565 NMBE 




1 


2 





CH 


BE 


Bern, Obstberg 


4 


9 


2000 


565 NMBE 


Balkon 





1 





CH 


BE 


Bern, Obstberg 


29 


10 


2000 


565 NMBE 


Wohnung 








4 


CH 


BE 


Bern, Obstberg 


27 


5 


2001 


565 NMBE 




1 








CH 


BE 


Bern, Obstberg 


2 


7 


2001 


565 NMBE 


Balkon 


1 


1 





1 CH 


BE 


Bern, Obstberg 


8 


7 


2001 


565 NMBE 


Haus 





1 





CH 


BE 


Bern, Obstberg 


25 


7 


2001 


565 NMBE 


Balkon 


4 


1 





CH 


BE 


Bern, Obstberg 


27 


7 


2001 


565 NMBE 


Balkon 


2 





1 


CH 


BE 


Bem, Obstberg 


31 


7 


2001 


565 NMBE 







1 





1 CH 


BE 


Bem, Obstberg 


8 


8 


2001 


565 NMBE 


Haus 


1 








CH 


BE 


Bem, Obstberg 


19 


8 


2001 


565 NMBE 




1 


1 





1 CH 


BE 


Bem, Obstberg 


21 


8 


2001 


565 NMBE 


Haus 





1 





CH 


BE 


Bern, Obstberg 


7 


9 


2001 


565 NMBE 







1 





1 CH 


BE 


Bern, Obstberg 


18 


10 


2001 


565 NMBE 


Haus 





1 





1 CH 


BE 


Bern, Obstberg 


21 


10 


2001 


565 NMBE 


Haus 


1 








CH 


BE 


Bern, Obstberg 


6 


9 


2002 


565 NMBE 


Haus 





1 





CH 


BE 


Bolligen 


7 


8 


2001 


580 NMBE 


Haus 


1 


2 





ü CH 


BE 


Burgdorf 






2002 


544 NMBE 







1 





CH 


BE 


Burgistein 


4 


10 


1999 


751 NMBE 







1 





CH 


BE 


Ittigen 


10 


8 


2000 


526 NMBE 


Haus 


1 








CH 


BE 


Leissigen 




7 


2000 


570 NMBE 


7.-8.2000 





1 





CH 


BE 


Münsingen 


27 


5 


2001 


531 NMBE 


Haus 


1 








CH 


BE 


Muri 


20 


6 


2000 


511 NMBE 


Badezimmer 


1 








CH 


BE 


Muri 


22 


6 


2000 


511 NMBE 


Badezimmer 





1 





CH 


BE 


Muri, Gümligen 


28 


6 


2000 


561 UGZ 


Haus 


1 








CH 


BE 


Thun 


25 


7 


1992 


560 NMBE 


Nähe der Aare 


1 








CH 


BE 


Worb 


26 


9 


2001 


581 NMBE 


Garten 


1 








CH 


BE 


Worb, Rüfenacht 


24 


6 


1998 


590 NMBE 







1 





CH 


BE 


Worb, Rüfenacht 


10 


9 


1999 


590 NMBE 




1 








CH 


BE 


Worb, Rüfenacht 


25 


9 


2001 


590 NMBE 


Garten 


1 








CH 


BE 


Worb, Rüfenacht 


22 


11 


2001 


590 NMBE 


Haus 


1 


3 


so 


2 CH 


BL 


Bottmingen, Bruderholz* 






1999 


365 NHMB 


August-Dezember, am Fenster 



420 



H. BAUR ET AL. 



Appendix 2 (Fortsetzung) 



e? 


9 


Ny 


Do Ld 


Kt 


Ort 


Tag 


Mon 


Jahr 


Höhe Coli. 


Bemerkungen 


1 








CH 


BL 


Füllinsdorf 




6 


2000 


330 NHMB 







1 





CH 


BL 


Liestal 




10 


2002 


450 NHMB 




1 








CH 


BL 


Münchenstein, Teufelsgraben 


8 


6 


2002 


270 NHMB 


Bachufer 


1 








CH 


BL 


Oberwil 






1997 


305 NHMB 


Sommer, Balkon (2. Stock) 


1 








CH 


BL 


Oberwil 






2002 


305 NHMB 


Juli-August, Wohnung 





1 





CH 


BL 


Sissach 




12 


1996 


370 NHMB 


Keller, Totfund 


1 








CH 


BL 


Sissach 


1 


4 


1999 


370 NHMB 


ex Nympha 30.6.1999 


1 








CH 


BS 


Basel 






195? 


270 NHMB 


Sammeljahr unsicher! 


1 








CH 


BS 


Basel 


5 


7 


2000 


270 NMBE 




3 


3 





CH 


BS 


Basel, Bruderholz 






1996 


310 NHMB 


Juli-August 





1 





CH 


BS 


Basel, Bruderholz 


25 


8 


1996 


310 NHMB 




3 








CH 


BS 


Basel, Bruderholz 


17 


9 


1999 


332 NHMB 


Haus 





1 





CH 


BS 


Basel, Bruderholz 




10 


1999 


332 NHMB 


Küche 





1 





CH 


BS 


Basel, Bruderholz 


1 


7 


2000 


355 NHMB 


Haus 





1 





1 CH 


BS 


Basel, Grossbasel west 




10 


2002 


276 NHMB 












CH 


BS 


Basel, Gundeldinger Quartier 


17 


9 


1995 


281 NHMB 












CH 


BS 


Basel, Gundeldinger Quartier 


7 


9 


1999 


281 NHMB 


Wohnung 





1 





CH 


BS 


Basel, Kleinbasel 


13 


9 


1999 


260 NHMB 


Schlafzimmer (2. Stock) 










CH 


BS 


Basel, Kleinbasel 


24 


10 


1999 


260 NHMB 


Wohnung 





1 





CH 


BS 


Basel, Kleinbasel, Horburg 


5 


8 


2001 


254 NHMB 


auf Fruchtstand von Daucus carota 










CH 


BS 


Basel, Kleinbasel, Lange Erlen 


8 


6 


2001 


260 NHMB 


ex Nympha 29.6.2001 










CH 


BS 


Basel, Kleinhüningen 


23 


6 


2000 


260 NHMB 


Badischer Rangierbahnhof 





1 





1 CH 


BS 


Riehen 






1999 


282 NHMB 


Oktober-November 


2 





1 


CH 


BS 


Riehen 


8 


6 


2000 


286 NHMB 


an Buxus sempervirens 





1 





CH 


BS 


Riehen 


4 


7 


2000 


282 NHMB 




4 


2 





CH 


BS 


Riehen 


26 


7 


2000 


285 NHMB 


am Fenster (Licht) 








3 


CH 


GE 


Lancy 


30 


8 


2000 


400 NMBE 


Storenkasten von Mietwohnungen 


1 








CH 


GR 


Grono 




7 


1887 


332 NMBE 




1 








CH 


GR 


Pontresina 






1900 


1800 CAN 




1 








CH 


GR 


Roveredo 


23 


7 


1933 


292 CAN 


23.7.-3.8.1933 


1 


1 





CH 


GR 


Soglio 




8 


1920 


1100 CAN 


Syntypen E. neolividus Fruhstorfer 





1 





CH 


GR 


Soglio 


21 


7 


1929 


1100 CKH 




2 








CH 


GR 


Soglio 


21 


7 


1929 


1100 CAN 










1 


CH 


GR 


Soglio 


21 


7 


1929 


1100 CAN 


Bestimmung unsicher 


1 








CH 


LU 


Buchrain 


24 


6 


2000 


459 NML 


auf Balkon 





1 





CH 


LU 


Emmen, Emmenbrücke 


21 


10 


1993 


436 NML 


Wohnung 





3 





CH 


LU 


Littau 


4 


8 


2000 


511 NML 


Balkon, Wohnung 


1 








CH 


LU 


Luzern 


20 


7 


1993 


430 NML 


Wohnung 





1 





CH 


LU 


Luzern 


8 


8 


1993 


430 NML 


Küche, Zimmer 


1 








CH 


LU 


Luzern 


12 


8 


1993 


430 NML 


Wohnung 


1 








CH 


LU 


Luzern 


19 


8 


1993 


430 NML 


Balkon 





1 





CH 


LU 


Luzern 


21 


8 


1993 


430 NML 


Küche, Schlafzimmer 


1 


1 





CH 


LU 


Luzern 


7 


9 


1993 


430 NML 


Balkon, Wohnung 





1 





CH 


LU 


Luzern 


7 


9 


1993 


430 NML 


Wohnung 





1 





CH 


LU 


Luzern 


10 


7 


2000 


430 NML 


Wohnung 


1 








CH 


LU 


Luzem 


12 


7 


2000 


430 NML 


Ausstellungsraum 


1 








CH 


LU 


Luzern 


2 


8 


2000 


430 NML 


Präparatorium 


1 


1 





CH 


LU 


Luzem 


9 


8 


2000 


430 NML 


Wohnung 


1 








CH 


LU 


Luzern 


28 


8 


2000 


430 NML 


Wohnung 


1 


1 


1 


CH 


LU 


Meggen 


17 


10 


1999 


440 NHMB 




1 








CH 


LU 


Meggen 




6 


2000 


440 NHMB 




2 








CH 


LU 


Rothenburg 


16 


7 


1993 


490 NML 


Wohnung 


2 


1 





CH 


LU 


Sursee 


19 


7 


2000 


504 NMBE 


Terrasse 


1 








CH 


LU 


Sursee 


22 


8 


2000 


504 NMBE 


Wohnung 





1 





CH 


LU 


Sursee 


25 


7 


2000 


504 NML 


Wohnung 


2 








CH 


NW 


Ennetbürgen 


27 


6 


2000 


440 NMBE 


Balkon 


1 


1 





CH 


ow 


Engelberg 


5 


8 


1993 


1000 NML 


Haus 





1 





CH 


ow 


Samen 




9 


1996 


470 NMBE 


Haus 








1 


CH 


SG 


Amden 


4 


1 


2001 


908 UGZ 


Ferienchalet, Fundjahr 2001 








1 


CH 


SG 


Jona 


3 


11 


2000 


433 UGZ 


Haus 


1 








CH 


SG 


Schmerikon 


22 


6 


2000 


408 UGZ 




1 


3 





CH 


SG 


Schmerikon 


21 


7 


2000 


408 UGZ 


Küche und Wohnzimmer 


2 


2 





CH 


SH 


Neuhausen 




11 


2001 


440 NMBE 


Haus 





1 





CH 


SH 


Schaffhausen 


24 


8 


2000 


550 UGZ 




1 


h 





CH 


SH 


Schaffhausen. Alter Emmersberg 


5 


6 


2000 


550 NMBE 





ECTOBIUS VimVENTRlS (BLATTODEA) 



421 



Appendix 2 (Fortsetzung) 



c? 


9 


\y 


Oo Ld 


Kt 


Ort 


Tag 


Mon 


Jahr 


Höhe Coll. 


Bemerkungen 





1 


1 


CH 


SO 


Riedholz 


10 


10 


2001 


530 NMBE 


Haus 


1 








CH 


SO 


Trimbach, Düriberg, Paradisli 


1 


7 


2001 


470 CGA 


Waldrand 





1 





CH 


SO 


Wangen bei Olten, Haftlet 


5 


10 


2001 


505 CGA 


Waldrand 





1 





CH 


sz 


Bach 


23 


8 


2000 


411 UGZ 


Bank 








1 


CH 


sz 


Freienbach, Pfäffikon 


8 


5 


2000 


412 UGZ 


Badezimmer 


1 


1 


1 


CH 


sz 


Wollerau 


31 


8 


2000 


504 UGZ 


Garten und Haus 





1 





CH 


TI 


Arzo 


20 


9 


1919 


500 ETHZ 


Arzo-Meride 


1 





1 


CH 


TI 


Arzo, Cava di marmo 


25 


6 


2001 


540 NMBE 


aufRubus 





2 





CH 


TI 


Arzo-Meride 


20 


9 


1919 


500 ZMB 







2 





CH 


TI 


Balerna 




9 


1919 


300 ETHZ 


S. Stefano-Balerna 





6 





CH 


TI 


Brusino-Arsizio 


12 


8 


1983 


274 MHNG 












CH 


TI 


Capolago 


7 


9 


1919 


274 ETHZ 


Capolago-Meride 










CH 


TI 


Caslano, Monte 


4 


9 


1971 


526 ETHZ 












CH 


TI 


Caslano, Monte 


9 


9 


1971 


526 ETHZ 












CH 


TI 


Cevio 


7 


10 


1981 


416 MHNG 












CH 


TI 


Chiasso, Pedrinate 




7 


1919 


450 ETHZ 




1 







CH 


TI 


Gandria, Cantine di Gandria 


13 


7 


1989 


290 CAN 


272-300m 










CH 


TI 


Generoso-Crocetta 


29 


7 


1919 


1360 ZMB 












CH 


TI 


Gordola 


13 


10 


1987 


350 ETHZ 




1 







CH 


TI 


Ligometto 




8 


1919 


500 CAN 












CH 


TI 


Ligometto 


28 


7 


1919 


500 ETHZ 




1 








CH 


TI 


Ligometto 


28 


7 


1919 


500 ZMB 







2 





CH 


TI 


Ligornetto-Meride 


17 


9 


1919 


500 ZMB 




1 


2 





CH 


TI 


Ligornetto-Tremona 




8 


1919 


500 ZMB 







1 





CH 


TI 


Mende 


17 


9 


1919 


580 ETHZ 


Ligornetto-Meride 





2 





CH 


TI 


Meride 


27 


9 


1999 


580 NMBE 




3 


1 


3 


CH 


TI 


Mende 


25 


6 


2001 


580 NMBE 







2 





CH 


TI 


Meride, Campagna 


25 


6 


2001 


620 NMBE 




6 


2 





CH 


TI 


Meride, San Antonio 


26 


6 


2001 


570 NMBE 







2 





CH 


TI 


Monte Boglia 


7 


9 


1961 


1050 CAN 












CH 


TI 


Monte Generoso 


12 


9 


1960 


CKH 


genaue Höhe unbekannt! 










CH 


TI 


Sonvico, Passo Pairolo 


20 


6 


1919 


1406 ZMB 




2 


2 





CH 


TI 


Tremona 




8 


1919 


575 ETHZ 


Ligometto-Tremona 










1 CH 


TI 


Villa Luganese, Denti della Vecchia 


5 


9 


1919 


1000 ZMB 


Syntypus E. neolividus Fruhstorfer 


1 








CH 


VD 


Chavannes-près-Renens 


16 


8 


1998 


403 UGZ 


Wohnung 


1 


2 





CH 


VD 


Chavannes-près-Renens 


20 


9 


2000 


403 UGZ 


Aussen, Terrasse 


2 








CH 


VD 


Jongny 


18 


8 


2000 


550 UGZ 


Aussen 










CH 


VD 


Lausanne 


20 


8 


1997 


500 UGZ 


Wohnung 










1 CH 


VD 


Lausanne 




8 


1998 


500 UGZ 


Aussen 








1 


CH 


VD 


Lausanne 


25 


4 


2000 


500 UGZ 


Wohnung 


1 







1 CH 


VD 


Lausanne 


10 


7 


2000 


500 UGZ 


Wohnung 










1 CH 


VD 


Lausanne 


2 


8 


2000 


500 UGZ 


Haus und Umgebung 


1 








CH 


VD 


Lausanne 


22 


8 


2000 


500 UGZ 


Haus 










CH 


VD 


Lausanne 


20 


11 


2000 


500 UGZ 


Kindergarten 










CH 


VD 


Montreux, Clarens 


14 


8 


1997 


400 UGZ 


Haus 





2 





CH 


VD 


Morges 


3 


10 


2000 


375 UGZ 


Spital, in Schabenfalle 


1 







CH 


VD 


Pully 


17 


8 


2000 


425 UGZ 


Haus 







1 


CH 


VD 


Pully 


21 


8 


2000 


425 UGZ 


Wohnung 







1 


CH 


VD 


Pully 


7 


9 


2000 


425 UGZ 


Wohnung 










CH 


VD 


Pully 


14 


9 


2000 


425 UGZ 


Wohnung 


1 







CH 


VD 


Tour-de-Peilz, La 


19 


7 


1996 


400 UGZ 


Wohnung 


1 








CH 


VD 


Vevey 


14 


8 


1998 


380 UGZ 


Wohnung 


1 








CH 


VD 


Veytaux 


10 


9 


2000 


403 UGZ 


Hotel 










CH 


vs 


Zwischbergen, Gondo 


9 


9 


1957 


840 ETHZ 






1 









CH 
CH 


vs 
vs 








1900 


ETHZ 
CAN 




1 








CH 


ZH 


Adliswil 


12 


10 


1999 


451 UGZ 


Wohnung 


3 







CH 


ZH 


Adliswil 


5 


7 


2000 


451 UGZ 




1 








CH 


ZH 


Affoltern aA. 


14 


10 


1999 


494 UGZ 


Wohnung 










CH 


ZH 


Affoltem aA. 


5 


9 


2000 


494 UGZ 


Haus und Umgebung 


1 








CH 


ZH 


Affoltern aA. 


2 


10 


2000 


494 UGZ 


Garten und an Fenstern 


4 








CH 


ZH 


Bonstetten 


7 


7 


2000 


528 UGZ 


Haus 










1 CH 


ZH 


Biilach 


6 


9 


2000 


420 UGZ 


Haus 


1 


2 


1 


CH 


ZH 


Biilach 








420 UGZ 


Wohnung 







'1 


CH 


ZH 


Dachsen 


25 


8 


2000 


393 UGZ 


Garten und Wohnung 



422 



H. BAUR ET AL. 



Appendix 2 (Fortsetzung) 



s 


5 


My 


Do Ld 


Kt 


Ort 


Tag 


Vlon 


Jahr 


Höhe Coli. 


Bemerkungen 










CH 


ZH 


Dällikon 


3 


7 


2000 


438 UGZ 


Wohnung 










CH 


ZH 


Dietlikon 


29 


6 


2000 


444 UGZ 


Wohnung und Gartensitzplatz 










CH 


ZH 


Dietlikon 


29 


6 


2000 


444 UGZ 


Restaurant 










CH 


ZH 


Dübendorf 


5 


7 


2000 


440 UGZ 


Attikawohnung 




1 





CH 


ZH 


Dübendorf 


5 


7 


2000 


440 UGZ 


Wohnung 


2 


1 





CH 


ZH 


Dübendorf 


6 


7 


2000 


440 UGZ 


Schlafzimmer 





1 





CH 


ZH 


Dübendorf 


6 


7 


2000 


440 UGZ 


Schlafzimmer 





1 





1 CH 


ZH 


Dürnten 


31 


8 


2000 


510 UGZ 







1 





CH 


ZH 


Egg 


2 


11 


1999 


540 UGZ 


Wohnung 


1 








CH 


ZH 


Embrach 


21 


6 


2000 


440 UGZ 


Haus 


1 








CH 


ZH 


Erlenbach ZH 


30 


6 


2000 


422 UGZ 


Wohnung 





2 





CH 


ZH 


Fällanden, Benglen 


15 


8 


2000 


600 UGZ 


Balkon und Wohnung 





1 





1 CH 


ZH 


Fällanden, Pfaffhausen 


15 


10 


1999 


600 UGZ 


Haus 





2 





CH 


ZH 


Geroldswil 


11 


7 


2000 


420 UGZ 


Kath. Pfarramt 





1 





CH 


ZH 


Geroldswil 


13 


7 


2000 


420 UGZ 


Haus 


2 








CH 


ZH 


Glattfelden 


18 


8 


2000 


410 UGZ 


Wohnung 





1 





CH 


ZH 


Greifensee 


19 


10 


1999 


440 UGZ 


Haus 








1 


CH 


ZH 


Hinwil, Wernetshausen 


7 


12 


1999 


800 UGZ 


Haus 





1 





CH 


ZH 


Hombrechtikon 


7 


7 


2000 


500 UGZ 


Haus 








3 


CH 


ZH 


Horgen 


12 


4 


2000 


410 UGZ 


Aussen, unter Blumentöpfen 





1 


4 


CH 


ZH 


Illnau 


6 


7 


2000 


510 UGZ 


Wohnung 





1 





CH 


ZH 


Illnau, Brunnacher, Ottikon 


12 


7 


2000 


600 UGZ 


Haus 





1 





CH 


ZH 


Küsnacht 


17 


10 


1961 


410 ETHZ 


Haus 


1 








CH 


ZH 


Küsnacht 


20 


12 


2000 


410 UGZ 


Haus 


1 








CH 


ZH 


Maur, Ebmatingen 


27 


7 


2000 


610 UGZ 


Wohnung 








1 


CH 


ZH 


Meilen, Feldmeilen 


6 


8 


2000 


480 UGZ 


Wohnung 








1 


CH 


ZH 


Meilen, Rossbachtobel 


19 


3 


2000 


470 UGZ 


Balkonkistchen 





1 


1 


CH 


ZH 


Neftenbach 


19 


9 


2000 


420 UGZ 


Haus 





1 





1 CH 


ZH 


Nürensdorf 


22 


10 


1999 


505 UGZ 


Haus 





1 





CH 


ZH 


Oberembrach 


25 


9 


1988 


460 ETHZ 







2 





CH 


ZH 


Oerwil am See 


11 


10 


1999 


538 UGZ 


Wohnung 





1 





CH 


ZH 


Oetwil am See 


4 


9 


2000 


538 UGZ 


Haus 





1 





CH 


ZH 


Opfikon 


18 


7 


2000 


455 UGZ 


Küche 





1 





CH 


ZH 


Otelfingen 


18 


10 


1999 


427 UGZ 


Wohnung 





2 





CH 


ZH 


Otelfingen 


18 


10 


1999 


427 UGZ 


Garten 


1 








CH 


ZH 


Otelfingen 


20 


6 


2000 


427 UGZ 


Treppenhaus 


1 








CH 


ZH 


Pfäffikon 


4 


8 


2000 


540 UGZ 


Wohnung 


1 








CH 


ZH 


Regensdorf 


4 


11 


1999 


460 UGZ 


Haus 





1 





CH 


ZH 


Richters wil 


17 


11 


1999 


410 UGZ 


Wohnung 





1 





CH 


ZH 


Richterswil 


14 


7 


2000 


410 UGZ 


Haus/Estrich 


1 


1 


1 


CH 


ZH 


Richters wil 


5 


7 


2000 


410 UGZ 







1 





CH 


ZH 


Schlieren 


2 


10 


2000 


390 UGZ 


Haus 





1 





CH 


ZH 


Seegräben, Aathal, Linggenberg 






2001 


510 UGZ 




1 








1 CH 


ZH 


Uitikon 


17 


7 


2000 


553 UGZ 


Haus 


1 








CH 


ZH 


Uitikon, Waldegg 


30 


6 


2000 


620 UGZ 


Bad/Küche 





1 





CH 


ZH 


Unterengstringen 


29 


9 


2000 


410 UGZ 


Wohnung 





1 





CH 


ZH 


Urdorf 


15 


10 


1999 


420 UGZ 


Küche 


1 








CH 


ZH 


Uster 


11 


9 


1990 


460 NMBE 


Haus 


1 








CH 


ZH 


Uster 


10 


10 


1993 


460 NMBE 


Haus 


1 


1 





1 CH 


ZH 


Uster 


23 


9 


1999 


460 NMBE 


Haus 





1 





CH 


ZH 


Uster 


9 


10 


1999 


460 NMBE 


Haus 





1 





CH 


ZH 


Uster 


31 


10 


1999 


460 NMBE 


Haus 


2 


1 


1 


CH 


ZH 


Uster 




10 


1999 


460 NMBE 


Haus 


1 








CH 


ZH 


Uster 


5 


6 


2000 


460 NMBE 


Haus 





1 


.0 


CH 


ZH 


Uster 


7 


7 


2000 


460 UGZ 


Haus 


1 


3 





1 CH 


ZH 


Uster 


19 


8 


2000 


460 NMBE 


Haus, 19.-23.8.2000 


1 


3 





1 CH 


ZH 


Uster 


23 


8 


2000 


460 NMBE 


Haus, 19.-23.8.2000 








3 


CH 


ZH 


Uster 


4 


9 


2000 


460 NMBE 


Haus 








3 


CH 


ZH 


Uster 


12 


9 


2000 


460 NMBE 


Haus 


1 








CH 


ZH 


Uster 


27 


9 


2000 


460 NMBE 


Haus 


2 


1 





CH 


ZH 


Uster 




10 


2000 


460 NMBE 


Haus 


1 








CH 


ZH 


Uster, Nänikon 


8 


8 


2000 


457 UGZ 


Wohnung 


2 


1 





CH 


ZH 


Volketswil 


15 


8 


2000 


465 UGZ 


Haus und Umgebung 





3 


1 


CH 


ZH 


Volketswil 


21 


8 


2000 


465 UGZ 







1 





CH 


ZH 


Wädenswil 


13 


10 


1999 


500 UGZ 


Haus 



ECTOBIUS VITTIVENTRIS (BLATTODEA) 



423 



Appendix 2 (Fortsetzung) 



tf 9 Ny Oo Ld Kt Ort 



Tag Mon Jahr Höhe Coll. Bemerkungen 



1 


3 


1 


1 CH 


ZH 


Wädenswil 


4 


3 


4 


1 CH 


ZH 


Wallisellen 





1 





CH 


ZH 


Weiningen, Fahrweid 





1 





CH 


ZH 


Wettswil am Albis 





1 





CH 


ZH 


Wetzikon 


2 








CH 


ZH 


Wetzikon 


1 








CH 


ZH 


Winterthur 


1 








CH 


ZH 


Winterthur 


1 


1 





CH 


ZH 


Winterthur 





1 





CH 


ZH 


Winterthur 








1 


CH 


ZH 


Winterthur 


1 








CH 


ZH 


Winterthur 


1 








CH 


ZH 


Winterthur 


1 








CH 


ZH 


Winterthur 





1 





CH 


ZH 


Winterthur 





1 





CH 


ZH 


Winterthur 





2 





1 CH 


ZH 


Winterthur 








1 


CH 


ZH 


Winterthur 


1 








CH 


ZH 


Winterthur 








1 


CH 


ZH 


Winterthur 








1 


CH 


ZH 


Winterthur 


1 


2 





CH 


ZH 


Winterthur 





1 





CH 


ZH 


Winterthur 


1 


1 





CH 


ZH 


Zollikon 





2 





CH 


ZH 


Zürich 





2 





CH 


ZH 


Zürich 


1 








CH 


ZH 


Zürich 


1 


3 





CH 


ZH 


Zürich 








1 


CH 


ZH 


Zürich 


1 


2 





CH 


ZH 


Zürich 





1 





CH 


ZH 


Zürich 


1 








CH 


ZH 


Zürich 





1 





CH 


ZH 


Zürich 








1 


CH 


ZH 


Zürich 


1 








CH 


ZH 


Zürich 








1 


CH 


ZH 


Zürich 








1 


CH 


ZH 


Zürich 








1 


CH 


ZH 


Zürich 








1 


CH 


ZH 


Zürich 


1 





3 


CH 


ZH 


Zürich 


2 








CH 


ZH 


Zürich 


2 


1 


1 


CH 


ZH 


Zürich 


1 








CH 


ZH 


Zürich 


1 








CH 


ZH 


Zürich 





1 


2 


CH 


ZH 


Zürich 


2 








CH 


ZH 


Zürich 





1 





CH 


ZH 


Zürich 


2 








CH 


ZH 


Zürich 


1 








CH 


ZH 


Zürich 


1 








CH 


ZH 


Zürich 


1 








CH 


ZH 


Zürich 


1 








CH 


ZH 


Zürich 


3 




1 


CH 


ZH 


Zürich 


3 







CH 


ZH 


Zürich 


1 







CH 


ZH 


Zürich 


1 







CH 


ZH 


Zürich 










CH 


ZH 


Zürich 










CH 


ZH 


Zürich 










CH 


ZH 


Zürich 


1 







CH 


ZH 


Zürich 


3 




1 


CH 


ZH 


Zürich 










CH 


ZH 


Zürich 


2 


2 


1 


CH 


ZH 


Zürich 


1 








CH 


ZH 


Zürich 





2 





CH 


ZH 


Zürich 



28 


9 


2000 


500 


UGZ 


Haus 


14 


9 


2000 


430 


UGZ 


Wohnung 


5 


7 


2000 


390 UGZ 


Küche 


21 


10 


1999 


562 


UGZ 


Haus 




10 


1999 


530 NHMB 


via "Deso Star" ans Museum gelangt 


5 


9 


2000 


530 UGZ 


Haus 


20 


8 


1997 


439 NMBE 


Haus 


18 


7 


1999 


439 NMBE 


Wohnung 


6 


8 


1999 


439 NMBE 


Wohnung 


28 


8 


1999 


439 NMBE 


Wohnung 


5 


9 


1999 


439 NMBE 


Wohnung 


5 


9 


1999 


439 NMBE 


Wohnung 


7 


9 


1999 


439 NMBE 


Wohnung 


9 


9 


1999 


439 NMBE 


Wohnung 


8 


10 


1999 


439 NMBE 


Hauseingang 


14 


10 


1999 


439 UGZ 


Haus 


15 


10 


1999 


439 UGZ 


Schlafzimmer 


24 


10 


1999 


439 NMBE 


Wohnung 


1 


11 


1999 


439 NMBE 


auf Prunus laurocerasus 


14 


12 


1999 


439 UGZ 


Haus 


17 


4 


2000 


439 NMBE 


Hauseingang 


2 


8 


2000 


439 


UGZ 


Haus und Umgebung 


9 


8 


2000 


439 UGZ 


Haus 


15 


10 


1999 


480 UGZ 


Büro 




8 


1992 


411 


ETHZ 




1 


8 


1995 


411 


ETHZ 




11 


9 


1995 


411 


ETHZ 




12 


10 


1996 


411 


NMBE 


Hauseingang 


17 


9 


1999 


411 


UGZ 


Haus 


17 


9 


1999 


411 


UGZ 


Haus 


5 


10 


1999 


411 


UGZ 


Wohnung 


15 


10 


1999 


411 


UGZ 


Küchenschrank 


28 


10 


1999 


411 


UGZ 


Wohnung 


11 


11 


1999 


411 


UGZ 


Haus 


11 


11 


1999 


411 


UGZ 


Haus 


19 


11 


1999 


411 


UGZ 


Geranien 


9 


3 


2000 


411 


UGZ 


Hausmauer 


5 


5 


2000 


411 


UGZ 


Wohnung 


16 


6 


2000 


411 


UGZ 


Haus 


22 


6 


2000 


411 


UGZ 




22 


6 


2000 


411 


UGZ 




26 


6 


2000 


411 


UGZ 


Haus und Umgebung 


27 


6 


2000 


411 


UGZ 


Küche 


7 


7 


2000 


411 


UGZ 


Haus 


10 


7 


2000 


411 


UGZ 


Haus 


17 


7 


2000 


411 


UGZ 


Waschküche 


17 


7 


2000 


411 


UGZ 




19 


7 


2000 


411 


UGZ 


Wohnung 


20 


7 


2000 


411 


UGZ 


Restaurant 


24 


7 


2000 


411 


UGZ 


Fenstersims 


24 


7 


2000 


411 


UGZ 


Wohnung 


25 


7 


2000 


411 


UGZ 


Wohnung 


28 


7 


2000 


411 


UGZ 


Wohnung und auf Balkon 


2 


8 


2000 


411 


UGZ 


Wohnung 


2 


8 


2000 


411 


UGZ 


Wohnung 


3 


8 


2000 


411 


UGZ 


Küche 


3 


8 


2000 


411 


UGZ 


Treppenhaus 


3 


8 


2000 


411 


UGZ 




7 


8 


2000 


411 


UGZ 


Wohnung 


8 


8 


2000 


411 


UGZ 


Wohnung 


9 


8 


2000 


411 


UGZ 


Haus 


9 


8 


2000 


411 


UGZ 


Haus 


10 


8 


2000 


411 


UGZ 


Wohnung 


10 


8 


2000 


411 


UGZ 


Wohnung 


15 


8 


2000 


411 


UGZ 


Haus 



424 



H. BAUR ETAL. 



Appendix 2 (Ende) 



<3 


9 


Ny 


Oo Ld 


Kt 


Ort 


Tag Mon 


Jahr 


Höhe Coll. 


Bemerkungen 










CH 


ZH 


Zürich 


16 


8 


2000 


411 


UGZ 


Haus 


6 







CH 


ZH 


Zürich 


16 


8 


2000 


411 


UGZ 


Wohnung 










CH 


ZH 


Zürich 


17 


8 


2000 


411 


UGZ 


Wohnung 





2 





CH 


ZH 


Zürich 


18 


8 


2000 


411 


UGZ 


Büffet der Badeanstalt (offen) 










CH 


ZH 


Zürich 


22 


8 


2000 


411 


UGZ 


Wohnung 





2 





CH 


ZH 


Zürich 


25 


8 


2000 


411 


UGZ 


Haus und Umgebung 










CH 


ZH 


Zürich 


25 


8 


2000 


411 


UGZ 


Wohnung 


2 








CH 


ZH 


Zürich 


25 


8 


2000 


411 


UGZ 












CH 


ZH 


Zürich 


28 


8 


2000 


411 


UGZ 


Wohnung 


1 







CH 


ZH 


Zürich 


31 


8 


2000 


411 


UGZ 


Haus 








1 


CH 


ZH 


Zürich 


31 


8 


2000 


411 


UGZ 












CH 


ZH 


Zürich 


5 


9 


2000 


411 


UGZ 


Wohnung 


1 


6 





1 CH 


ZH 


Zürich 


13 


9 


2000 


411 


UGZ 


Haus 










CH 


ZH 


Zürich 


20 


9 


2000 


411 


UGZ 


Haus 





2 





1 CH 


ZH 


Zürich 


21 


9 


2000 


411 


UGZ 


Haus 










CH 


ZH 


Zürich 


27 


9 


2000 


411 


UGZ 












CH 


ZH 


Zürich 


28 


9 


2000 


411 


UGZ 


Hotel 


1 





1 


CH 


ZH 


Zürich 


20 





2000 


411 


UGZ 


Wohnung 








1 


CH 


ZH 


Zürich 


25 





2000 


411 


UGZ 


Wohnung, Toilette 










1 CH 


ZH 


Zürich 


26 





2000 


411 


UGZ 


Küche 










CH 


ZH 


Zürich 


26 





2000 


411 


UGZ 


Wohnung 


1 








CH 


ZH 


Zürich 


3 


1 


2000 


411 


UGZ 


Haus 










CH 


ZH 


Zürich 


8 


1 


2000 


411 


UGZ 


Haus 





4 





CH 


ZH 


Zürich 


13 


1 


2000 


411 


UGZ 


Haus 










CH 


ZH 


Zürich 


14 


1 


2000 


411 


UGZ 


Küche 








1 


CH 


ZH 


Zürich 


28 


1 


2000 


411 


UGZ 


Küche 










1 CH 


ZH 


Zürich 


29 


1 


2000 


411 


UGZ 


Haus 








1 


CH 


ZH 


Zürich, Affoltem 


24 


5 


2000 


460 UGZ 


Wohnzimmer und Küche 








1 


CH 


ZH 


Zürich, Albisrieden 


13 


4 


2000 


450 UGZ 


Balkon 








1 


CH 


ZH 


Zürich, Albisrieden 


24 


4 


2000 


450 UGZ 


Balkon 


1 








CH 


ZH 


Zürich, Altstetten 


8 


6 


2000 


440 


UGZ 


Küche 








1 


CH 


ZH 


Zürich, Altstetten 


16 


6 


2000 


440 UGZ 


Balkon 


1 








CH 


ZH 


Zürich, Altstetten 


21 


6 


2000 


440 


UGZ 


Haus 


1 








CH 


ZH 


Zürich, Altstetten 


22 


6 


2000 


440 UGZ 


Schlafzimmer 








1 


CH 


ZH 


Zürich, Höngg 


17 





2000 


500 UGZ 


Wohnung, Küche 


2 








CH 


ZH 


Zürich, Oerlikon 


20 


9 


1999 


465 


UGZ 


Haus 


2 








D 


BW 


Bad Krozingen 




8 


2002 




NHMB 


Wohnhaus 











D 


BW 


Gottenheim, NW Freiburg i.Br. 






2002 




NHMB 


Siedlung; 3 Ex. (keine Belege) 


1 








D 


BW 


Lörrach 




7 


2002 




NHMB 


Wohnhaus 





1 


1 


D 


BW 


Weil am Rhein 







2001 


270 NHMB 


Tierhandlung, gehäuftes Auftreten 


1 








I 


NI 


Bellagio, S.S. Giovanni/Como 


19 


7 


1981 




CAN 







2 





I 


NI 


Chiavenna, SSE Grat M. Matec 


1 


9 


1971 


1000 CAN 


950-1050 m 


1 








I 


NI 


Dorio, Ronchi Vesgallo/Como 


12 


8 


1963 


350 CAN 


250-450 m 


1 








I 


NI 


Mendotica/Ligurien 


14 


8 


1986 


880 CAN 







1 





I 


NI 


Pamparato, S.Grée/Piemonte 


2 


9 


1985 


1125 


CAN 


1100-1 150 m 


1 


2 





I 


NI 


Portese-Salò/Gardasee 


22 


8 


1921 




ZMB 


Syntypen E. vitreus Ramme 





1 





I 


NI 


Savogno/Friaul 


26 


8 


1973 


350 CAN 


200-500 m 


1 








I 


NI 


Sondrio, Albonico, Il Forno 


22 


7 


1981 


575 CAN 


550-600 m 


1 








I 


NI 


Sondrio, Olgiasca-Piona 


21 


7 


1981 


311 


CAN 


250-372 m 


1 








I 


NI 


Vatgumnu 


18 


7 


1959 




CAN 


Sumpfwiese 





1 





I 


NI 


Vestreno, Mad. Bondo/Como 


6 


9 


1963 


600 CAN 




29 








I 


IO 


S. Gimignano, 4,5km ESE 


4 


9 


1997 


130 NMBE 





Revue suisse de Zoologie 111 (2): 425-432; juin 2004 



Amblyrhynchichthys micr acanthus, a new species of cyprinid fish 
from Indochina (Cyprinif ormes: Cyprinidae) 

Heok Hee NG 1 & Maurice KOTTELAT 2 

iFish Division, Museum of Zoology, University of Michigan, 1109 Geddes Avenue, 

Ann Arbor, Michigan 48109-1079, USA (email: heokheen@umich.edu) 

Phone: 1-734-647-2192. Fax: 1-734-763-4080. 
2 Case postale 57, Cornol, CH-2952, Switzerland (email: mkottelat@dplanet.ch) 



Amblyrhynchichthys micr acanthus, a new species of cyprinid fish from 
Indochina (Cyprinif ormes: Cyprinidae). - Amblyrhynchichthys micra- 
canthus sp. n. is described from the Mekong, Chao Phraya, Mae Khlong and 
Tapi river drainages in Indochina (mainland Southeast Asia). It can be dis- 
tinguished from its only congener, A. truncatus, in having a more truncate 
snout (with a straight vs. gently rounded lateral profile), shorter dorsal spine 
(15.5-25.7 %SL vs. 26.7-29.9), smaller eye when similar-sized individuals 
of larger than ca. 70 mm SL are compared (28.7-34.9 %HL vs. 32.7-36.3), 
and fewer gill rakers (34-37 vs. 39-^-0). 

Key-words: Amblyrhynchichthys - Cyprinidae - new species - Indochina. 

INTRODUCTION 

Fishes of the cyprinid genus Amblyrhynchichthys Bleeker, 1860 (type species 
Barbus truncatus Bleeker, 1851) are easily distinguished from other cyprinids in 
having an extremely blunt snout, anteriorly situated nostrils, last simple dorsal-fin ray 
stout and serrated posteriorly, and a well-developed hyaline eyelid. To date, the genus 
has been considered monotypic, being found in river drainages throughout Sumatra, 
Borneo and Indochina (mainland Southeast Asia). The other nominal species referred 
to Amblyrhynchichthys, A. altus Vaillant, 1893, is considered a junior synonym of 
Kalimantania lawak (Bleeker, 1855) (see Bänärescu, 1980). 

A comparison of Indochinese, Sumatran and Bornean specimens previously 
identified as A. truncatus shows that two distinct species can be recognised, with the 
Indochinese one described herein as Amblyrhynchichthys micracanthus. In previous 
literature on Indochinese fishes (e.g. Roberts & Warren, 1994: 100; Rainboth, 1996: 86, 
PI. VII Fig. 50; Kottelat, 2001: 38, Fig. 25), A. micracanthus has been identified as A. 
truncatus (type locality: Banjarmasin in southern Borneo). The situation in 
Amblyrhynchichthys is one more example of a species once considered to have a very 
wide distribution in large rivers throughout Southeast Asia and now found to consist of 
two distinct species: one Sundaic (Borneo, Java, Sumatra and Malay Peninsula) and 
one northern Indochinese (Mekong and Chao Phraya basins). Other cases reported in 



Manuscript accepted 08.12.2003 



426 H. H. NG & M. KOTTELAT 

recent years include: Belodontichthys dinema and B. truncatus, Laides hexanema and 
L. longibarbis, Bagrichthys macropterus and B. obscurus, and Helicophagus waan- 
dersii and H. leptorhynchus (Kottelat & Ng, 1999; Ng 1999a, b; Ng & Kottelat, 2000) 
and we are aware of numerous other cases awaiting re-analysis. A model of the 
historical biogeography of species with this pattern of distribution has been proposed 
by Bornbusch & Lundberg (1989), who hypothesized that the post-Pleistocene 
isolation of the North Sunda River system resulted in speciation. 

MATERIAL AND METHODS 

Measurements were made point to point with dial callipers and data recorded to 
0. 1 mm. Counts and measurements were made on the left side of specimens whenever 
possible. Subunits of the head are presented as proportions of head length (HL). Head 
length itself and measurements of body parts are given as proportions of standard 
length (SL). Measurements and counts were made following Hubbs & Lagler (1947), 
except for scale counts, which follow those of Kottelat (1984, 1990), and pharyngeal 
tooth counts, which follow those of Jenkins & Burkhead (1994), but exclude the 
replacement teeth. 

Fin rays were counted under a binocular dissecting microscope using trans- 
mitted light. Material for this study is deposited in the following collections: California 
Academy of Sciences, San Francisco (CAS), collection of the second author, Cornol 
(CMK), Museum of Comparative Zoology, Cambridge (MCZ), Nationaal Natuur- 
historisch Museum, Leiden (RMNH), Museum of Zoology, University of Michigan, 
Ann Arbor (UMMZ), and the Zoological Reference Collection of the Raffles Museum 
of Biodiversity Research, Singapore (ZRC). 

DESCRIPTION 

Amblyrhynchichthys micracanthus sp. n. Figs 1 and 2a 

Holotype. UMMZ 232203, 100.2 mm SL; Cambodia: Kandal province, Tonle Sap River 
22 km upstream from Phnom Penh; W. J. Rainboth and N. van Zalinga, 16 January 1995. 

Paratypes. CAMBODIA: CAS 94193, 1 ex., 111.8 mm SL; market at Phnom Penh; T. 
R. Roberts and K. E. Witte, 18 January-20 February 1994. CAS 94285, 5 ex., 98.0-125.3 mm 
SL; Cambodia: Stung Treng; T. R. Roberts, 2-18 February 1994. UMMZ 181245, 2 ex., 97.0- 
122.2 mm SL; Pursat province, Great Lake at Kompong Luong; J. Bardach, 18 March 1959. 
UMMZ 232277, 1 ex., 103.3 mm SL; Stung Treng morning market; W. J. Rainboth, N. van 
Zalinga and C. Rotha, 26 January 1995. UMMZ 232527, 2 ex., 101.6-110.2 mm SL; Kandal 
province, Tonle Sap; W. J. Rainboth and C. Rotha, 13 February 1995. UMMZ 232622, 1 ex., 
68.5 mm SL; Kandal province, Prek Mong Ya, just upstream from confluence with Bassac River 
at fishing lot 10; W. J. Rainboth and C. Rotha. 20 February 1995. UMMZ 232713, 1 ex., 91.5 
mm SL; Kompong Thom province, Tonle Sap at exit to Great Lake, 4 km NW of Chhnok Trou, 
at Kompong Thom fishing lot 2; W. J. Rainboth, N. van Zalinga and C. Rotha, 28 February 1995. 
UMMZ 234374, 1 ex., 97.2 mm SL; Kandal province, Tonle Sap at Dai fishery row 9, 25 km 
upstream from Phnom Penh; W. J. Rainboth et al, 22 January 1996. UMMZ 235499, 3 ex., 79.3- 
91.5 mm SL; Kandal province. Tonle Sap at Dai fishery row 9, 15 km upstream from Phnom 
Penh; W. J. Rainboth et al, 27 January 1996. LAOS: CAS 94272, 1 ex., 102.4 mm SL; 
Champasak province, Mekong river at Ban Hang Khone, just below Khone Falls; T. R. Roberts, 
June-July 1993. CMK 13137, 6 ex., 59.4-68.1 mm SL; Vientiane province, Mekong River at 
mouth of Nam Mang and lower 100 m of Nam Mang, 18°21'48"N 103°14'16"E; M. Kottelat et 
al, 22 February 1997. UMMZ 235320, 2 ex., 135.8-170.6 mm SL; Champasak province, 



AMBLYRHYNCHICHTHYS MICRACANTHUS 427 




Fig. 1 
Amblyrhynchichthys micr acanthus, CMK 13137, paratype, 68.1 mm SL. 

Mekong River at Ban Hang Khone, just downstream from Khone Falls; I. G. Baird, date un- 
known. THAILAND: CAS 61877, 2 ex., 128.1-141.4 mm SL; Ubon Ratchathani province, fish 
market at Ubon Ratchathani; T. R. Roberts, 28 June 1985. CAS 93243, 1 ex., 95.8 mm SL; 
Ayutthaya province, Ayutthaya market; T. R. Roberts, 5-7 March 1989. CAS 93921, 2 ex., 95.9- 
144.7 mm SL; Nakhon Sawan province, Nakhon Sawan market; T. R. Roberts, 4-5 February 
1989. MCZ 47307, 165.2 mm SL; Mekong River at Nong Khai; T. R. Roberts, May 1970. 
UMMZ 195088, 1 ex., 125.1 mm SL; Nakhon Sawan province, Chao Phraya River, floodwaters 
20 km N of Nakhon Sawan; K. F. Lagler, 15 October 1964. UMMZ 195364, 39 mm SL; Maharaj 
province, Chao Phraya River, 17.5 km N of Ayutthaya; S. Mekbahn, 27 December 1964. UMMZ 
195856, 1 ex., 98.5 mm SL; Mae Nam Khwae Yai about 10 km upstream from Kanchanaburi; 
K. F Lagler, 24 March 1965. UMMZ 195883, 5 ex., 84.3-168.1 mm SL; Mae Nam Mae Khlong 
at 2 km downstream of Ban Pong; K. F Lagler et al., 25 March 1965. 

Non-types. CAMBODIA: UMMZ 234483, 2 ex., 36.1-51.2 mm SL; Kandal province, 
floodplain lake near Mekong. THAILAND: UMMZ 185226, 1 ex., 94.3 mm SL; vicinity of 
Bangkok. UMMZ 195396, 17 ex., 35.3-53.3 mm SL; Maharaj province, Kok Tong Canal (trib- 
utary of Chao Phraya River) 17.5 km N of Ayuthaya. UMMZ 195722, 4 ex., 62.9-73.5 mm SL; 
Ubon Ratchathani province, Mun River, about 20 km downstream from Ubon Ratchathani. 
UMMZ 195739, 1 ex., 64.1 mm SL; Ubon Ratchathani province, Mun River, about 5 km down- 
stream from Ubon Ratchathani. UMMZ 224265, 1 ex., 101.3 mm SL; Ubon Ratchathani 
province, Khong Chiam district, Mun River at Ban Dan, 1.2 km upstream from confluence with 
Mekong River. UMMZ 224296, 1 ex., 114.7 mm SL; Nakhon Phanom province, Mekong River, 
island off Ban Tha Kai, 20 km downstream from Mukdahan, 1 km from Thai, 5 km from Laos 
side. UMMZ 224311, 1 ex., 111.5 mm SL; UMMZ 224353, 1 ex., 128.4 mm SL; UMMZ 
224364, 1 ex., 137.7 mm SL; UMMZ 224391, 1 ex., 132.9 mm SL; UMMZ 224398, 1 ex., 121.6 
mm SL; UMMZ 224448, 2 ex., 114.7-196.7 mm SL; UMMZ 224455, 1 ex., 111.5 mm SL; Ubon 
Ratchathani province, Khong Chiam district, Mun River at Ban Dan, 3 km upstream from 
confluence with Mekong River. UMMZ 224508, 1 ex., 95.8 mm SL; Ubon Ratchathani province, 
stream from Bung Klang Huen to Mun River, 10 km from Mun River, 3 km E of Ubon 
Ratchathani. UMMZ 233820, 1 ex., 43.0 mm SL; Ubon Ratchathani province, Huay Mark, 8 km 
N of Khong Chiam at confluence with Mekong River. UMMZ 237228, 30 ex., 36.2-58.7 mm SL; 
Maharaj province, Kok Tong, 15 km N of Ayutthaya, tributary of Lop Buri and Chao Phraya 
rivers. VIETNAM: UMMZ 218147, 13 ex., 29.0-59.1 mm SL; Phong Dinh province, Bassac 



428 



H. H. NG & M. KOTTELAT 





FlG. 2 
Lateral views of heads of: A. Amblyrhynchichthys micr -acanthus, UMMZ 235320, paratype, 
135.8 mm SL; B. A. truncatus, CMK 4763, 137.8 mm SL. Scale bar indicates 5 mm. 



River 0.5 km downstream from Can Tho. UMMZ 218533, 9 ex., 27.2-47.2 mm SL; UMMZ 
218547, 19 ex., 31.0-48.8 mm SL; UMMZ 218568, 11 ex., 26.5-43.1 mm SL; UMMZ 218583, 
1 ex., 49.5 mm SL; Phong Dinh province, Bassac River at Can Tho. UMMZ 218675, 8 ex., 40.8- 
73.0 mm SL; Phong Dinh province, Bassac River adjacent to Dong Phu. UMMZ 218651, 91 ex., 
24.0-62.2 mm SL; An Giang province, Bassac River 1.3 km S of Long Xuyen. UMMZ 224853, 
1 ex., 49.5 mm SL; An Giang province, rice paddy 1 km S of Highway 10 in Hoa Binh Thanh 
district. UMMZ 224956, 1 ex., 97.5 mm SL; Phong Dinh province, Can Tho fish market. 

Diagnosis. Amblyrhynchichthys micracanthus can be distinguished from its 
sole congener, A. truncatus, in having a more truncate snout (with a straight vs. gently 
rounded lateral profile), shorter dorsal spine (15.5-25.7 %SL vs. 26.7-29.9), smaller 
eye when similar-sized individuals of larger than ca. 70 mm SL are compared (28.7- 
34.9 %HL vs. 32.7-36.3), and fewer gill rakers (34-37 vs. 39-40). 

Description. Morphometric and meristic data are given in Table 1. Body 
moderately deep and strongly compressed, with narrow predorsal midline. Predorsal 
profile smoothly rounded, with small concavity at nape and then sloping gently ven- 
trally from origin of dorsal fin to end of caudal peduncle. Ventral profile less convex 
than dorsal profile, sloping gently ventrally to anterior end of anal-fin base, then 
sloping more steeply dorsally from there to end of caudal peduncle. Lateral line 
complete. Scales with strongly convergent radii. 

Head narrow, snout extremely blunt, with a vertical lateral profile. Nostrils large 
and anteriorly situated. Eye in upper half of head, visible from dorsal and ventral 
aspects; moderately large. Hyaline eyelid well developed, covering anterior and 
posterior quarters of orbit. Suborbital bones fairly large, narrowest directly below eye. 
Mouth subterminal, with well-developed groove separating lips from both upper and 
lower jaws. Lips thick; lower lip present medially and with complete post-labial 
groove. Gill membranes broadly joined to isthmus. Gill rakers long, one third as long 
as opposing filaments on epibranchial and half as long as opposing filaments on 
ceratobranchial. Pharyngeal bones stout, with three rows (2,3,4-4,3,2) of curved, 



AMBLYRHYNCHICHTHYS MICRACANTHUS 



429 



Table 1. Morphometric and meristic data for Amblyrhynchichthys micracanthus (n=30: 35.7- 
170.6 mm SL) 



MORPHOMETRICS 

In%SL 

Head length 23.5-28.6 

Head width 12.0-16.4 

Head depth 16.9-20.6 

Body depth at dorsal origin 24.4-36.3 

Predorsal length 46.8-50.9 

Preanal length 70.6-77.2 

Prepelvic length 43.9^9.6 

Prepectoral length 23.5-27.5 

Length of dorsal-fin base 14.6-19.1 

Dorsal-spine length 15.5-25.7 

Anal-fin length 9.2-11.4 

Pelvic-fin length 17.6-21.0 

Pectoral-fin length 18.2-20.1 

Caudal-fin length 24.2-31.2 

Length of caudal peduncle 16.2-21.0 

Depth of caudal peduncle 7.6-13.0 
In %HL 

Snout length 22.0-28.0 

Interorbital distance 2 1 . 1-39.9 

Eye diameter 28.7-34.9 



MERISTICS 

Dorsal-fin rays 
Anal-fin rays 
Pelvic-fin rays 
Pectoral-fin rays 
Caudal-fin rays 
Lateral line scales 
Predorsal scales 
Scales in transverse lines 
Circumpeduncular scales 
Rakers on first gill arch 



Vertebrae 



iv,9 (30) 
iii,5 (30) 
i,8 (30) 
i,14 (30) 
i,9,8,i (30) 
35+2 (30) 
11 (30) 

145/1/3V5 (30) 
VüllßVi (30) 
10+24 (9), 
11+23(3), 
10+25 (7), 
11+24(3), 
10+26 (3), 
11+25 (3) or 
10+27 (2) 
20+12=32 (4) or 
20+13=33 (26) 



chisel-shaped teeth. Grinding surfaces of distalmost major teeth widest, width pro- 
gressively decreasing proximally. 

Dorsal fin moderately high, with concave distal margin, sharply pointed at apex; 
origin slightly anterior to pelvic-fin origin. Last unbranched ray longest; spinous part 
with 11 to 18 serrations on posterior edge. Pectoral fin long and falcate, extending to 
scale row at pelvic-fin origin; distal margin concave near tip but straight otherwise. 
Pelvic fin moderately falcate, extending to slightly more than midway between pelvic- 
fin origin and anterior base of anal-fin; distal margin nearly straight. Anal fin with 
strongly concave distal margin and last three posterior rays equal in length; when 
adpressed against body, extending to middle of caudal peduncle. Two scale rows 
between urogenital opening and anal-fin origin. Caudal peduncle strongly compressed 
and moderately long. Caudal fin deeply forked, upper and lower lobes pointed. 

Sexual dimorphism absent. Breeding tubercles absent and body depth between 
sexes not different. 

Colour. Preserved material with a pale brown body and a faint broad stripe 
consisting of aggregations of melanophores on sides of body immediately above lateral 
line. Dorsal and caudal fins hyaline, with a faint black posterior margin. All other fins 
hyaline. Colour in life an overall silvery white. 

Distribution. Lower and middle Mekong, Chao Phraya, Mae Khlong and Tapi 
river drainages in Indochina (mainland Southeast Asia). In the Mekong, it reaches 
upriver to Vientiane. 

Etymology. From the Greek mikros, meaning small, and akantha, meaning 
thorn. In reference to the smaller dorsal spine of this species compared to A. truncatus. 
Treated as an adjective. 



430 



H. H. NG & M. KOTTELAT 



Ecology. Amblyrhynchichthys micracanthus is found mainly in rivers, with 
juveniles occasionally entering swamps and flooded fields (Taki, 1978). In the Mekong 
River drainage, this species moves into the inundated forest during the flood season 
(June-September) and return to the river in October and November (Rainboth, 1996). 
It feeds mainly on periphyton; other minor food items include phytoplankton, 
zooplankton and benthic algae (Rainboth, 1996). 

DISCUSSION 

The difference in eye diameter between A. micracanthus and A. truncatus is 
only apparent when similar-sized individuals are compared. We were able to examine 
only two specimens smaller than 90 mm SL for A. truncatus, but when eye diameter is 
plotted against SL for both species (Fig. 3), the graph indicates that the comparison is 
only meaningful when specimens of the same size (larger than ca. 70 mm SL) are used. 



Q 

üJ 




a A. micracanthus 
• A. truncatus 



230 



SL (mm) 



Fig. 3 
Eye diameter (ED) plotted against standard length for Amblyrhynchichthys species. 

The morphometric differences observed between A. micracanthus and A. trun- 
catus cannot be explained by ontogeny alone. The regression lines in the biplots of eye 
diameter (Fig. 3) and dorsal-spine length (Fig. 4) against SL show are significantly 
different (ANCO VA. P=0.00111 and P<0.0000005 respectively). 

Although the different snout shapes of A. truncatus and A. micracanthus would 
suggest that the shapes of the maxilla, premaxilla and possibly the circumorbital bones 
would differ, no significant differences in the osteology of the two species were 
observed, suggesting that the differences in shape are more likely caused by soft tissue. 



AMBLYRHYNCHICHTHYS MICRACANTHUS 



431 




* A. micracanthus 

• A. truncatus 



230 



SL (mm) 



Fig. 4 
Dorsal-spine length (DSL) plotted against standard length for Amblyrhynchichthys species. 



Bleeker (1851) described A. truncatus on the basis of three specimens 50- 
110 mm TL [total length] from Banjarmasin, Borneo. We have examined 2 specimens 
from Bleeker's collection which were registered as syntypes. RMNH 9091 is 35.3 mm 
SL, which corresponds to a total length of 50 mm and therefore seems to be the 50 mm 
TL syntype. RMNH 7032 is 56.5 mm SL and is probably the third specimen, for which 
Bleeker did not indicate the size (between 50 and 110 mm TL). Other Bleeker speci- 
mens of A. truncatus in RMNH are too large to be syntypes (RMNH 17176, 4, 96.5- 
203.4 mm SL; the 96.5 mm SL specimen would have been at least 120 mm TL). The 
whereabouts of the third syntype are not known. The 'cotype' listed by Bertin & Estève 
(1948) cannot have a type status as it is too large and is from Sumatra. The two 
syntypes agree with A. truncatus as diagnosed above and RMNH 9091 is hereby 
designated as lectotype. 

COMPARATIVE MATERIAL 

Amblyrhynchichthys truncatus: BORNEO: RMNH 7032, paralectotype, 56.5 mm SL; 
RMNH 9091, lectotype, 35.3 mm SL; Kalimantan Selatan, Banjarmasin. CMK 10190, 2 ex., 
116.1-157.3 mm SL; Kalimantan Barat, Danau Genting, between Danau Pengembung and 
Danau Belida. UMMZ 209912, 2 ex., 95.5-115.4 mm SL; Kalimantan Barat, Sungai Tawang 
near Danau Pengembung. ZRC 40025, 1 ex., 146.5 mm SL; Kalimantan Selatan, Banjarmasin, 
Pasar Lima Beton. SUMATRA: CMK 4763, 2 ex., 133.2-137.8 mm SL; ZRC 38556, 1 ex., 147.0 
mm SL; ZRC 43160, 2 ex., 132.4-140.0 mm SL; ZRC 44109, 2 ex., 128.5-139.5 mm SL; Jambi, 
Pasar Angso Duo. MALAY PENINSULA: ZRC 1764, 1 ex., 175.8 mm SL; Malaysia: Pahang, 
King George V National Park. ZRC 1765, 1 ex., 173.0 mm SL; Malaysia: Pahang, Kuala Tahan. 



432 H. H. NG & M. KOTTELAT 



ACKNOWLEDGEMENTS 

We thank David Catania (CAS), Karsten Hartel (MCZ), Martien van Oijen 
(RMNH), Douglas Nelson (UMMZ) and Kelvin Lim (ZRC) for permission to examine 
material under their care. Funding from a Block Grant from the Department of Ecology 
and Evolutionary Biology, University of Michigan and the Rackham School of 
Graduate Studies, University of Michigan to the first author provided financial support 
for this project. 

REFERENCES 

Bänarescu, P. 1980. Kalimantania and Neobarynotus, two new Indonesian genera of minnows 

(Pisces, Cyprinidae). Travaux du Muséum d'Histoire Naturelle "Grigore Antipa" 22: 

471-478. 
Bertin, L. & Estève, R. 1948. Catalogue des types de poissons du Muséum National d'Histoire 

Naturelle. 4e partie. Ostariophysaires (Cyprinif ormes). Muséum National d'Histoire 

Naturelle, Paris, 117 pp. 

Bleeker, P. 1851. Bijdrage tot de kennis der ichthyologische fauna van Borneo, met beschrij- 
ving van 16 nieuwe soorten van zoetwatervisschen. Natuurkundige Tijdschrift voor 
Nederlandsch Indie 1: 1-16. 

Bornbusch A. H. & Lundberg, J. G. 1989. A new species of Hemisilurus (Siluriformes, 
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Jenkins, R. E. & Burkhead, N. M. 1994. Freshwater fishes of Virginia. American Fisheries 
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Kottelat, M. 1984. A new Rasbora s. 1. (Pisces: Cyprinidae) from northern Thailand. Revue su- 
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(Teleostei: Bagridae). Revista de Biologia Tropical 47: 545-552. 

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Revue suisse de Zoologie 111 (2): 433-456; juin 2004 



The centipedes of the Maltese Archipelago (Chilopoda) 

Marzio ZAPPAROLI 1 , Alessandro MINELLI 2 & Patrick J. SCHEMBRI 3 

1 Dipartimento di Protezione delle Piante, Università degli Studi della Tuscia, 
Via San Camillo de Lellis, 1-01100 Viterbo, Italy. 

2 Dipartimento di Biologia, Università di Padova, Via Ugo Bassi 58 B, 
1-35131 Padova, Italy. 

3 Department of Biology, University of Malta, Msida MSD06, Malta. 



The centipedes of the Maltese Archipelago (Chilopoda). - The chilopod 
fauna of the Maltese Islands (Malta, Gozo, Cornino) was studied from a 
faunistic and Zoogeographie point of view. A list of the species found on 
these islands is given, based on recent faunistic investigations as well as on 
a critical assessment of the few records available in the literature. Twenty- 
one species are recorded to occur on the islands: 1 Scutigeromorpha, 7 
Lithobiomorpha, 3 Scolopendromorpha, and 10 Geophilomorpha. Twenty 
species are confirmed to occur on the island of Malta, 1 1 on Gozo and three 
on Cornino. The Maltese chilopod fauna mostly consists of species that are 
widespread in the Mediterranean islands, particularly those of the western 
Mediterranean. As shown in other studies on the chilopod faunas of other 
Mediterranean micro-insular systems, that of the Maltese Islands is mainly 
influenced by ecological factors rather than by paleogeographic and paleo- 
climatic ones. Zoogeographically the Maltese chilopod fauna is mainly 
Mediterranean in character, with a very limited representation of Holarctic 
(22%) and European (11%) species. 

Key-words: Chilopoda - Maltese Islands - Malta - Gozo - Cornino - fauna - 
biogeography. 

INTRODUCTION 

Very few papers have been devoted to the Maltese centipedes so far. The first 
lists of species were published at the end of the nineteenth century by the Maltese 
naturalist Giovanni Gulia (Gulia, 1890, 1913), who recorded only three species, all 
probably collected and studied by himself. About eighty years later, Matic et al. (1967) 
listed another seven species from material collected in 1965 by Marcello La Greca and 
co-workers, as part of a research programme on the Mediterranean fauna carried out 
by the Zoological Institute of the University of Catania (Italy). After that, the only 
published work on this subject known to us is a list of 12 species from the Maltese 
Islands, in a study on the centipede fauna of the West Mediterranean area by Foddai 
et al. (1996). 



Manuscript accepted 07.11.2003 



434 M. ZAPPAROLI ET AL. 

Recently (mainly in the eighties), research on the Maltese fauna has been taken 
up by workers from the University of Malta and ample material has been collected 
from the main islands of the archipelago. This material, together with specimens occa- 
sionally collected by Italian universities and natural history museums, is the subject of 
the present study. 

The aim of this paper is to list and discuss this material and all the previously 
published records of chilopods from the Maltese islands. 

STUDY AREA 

The Maltese archipelago (Fig. 1), situated in the central Mediterranean, appro- 
ximately 96 km from Sicily and 290 km from North Africa, consists of three inhabited 
islands, i.e. Malta (246.5 km 2 ), Gozo (65.8 km 2 ) and Cornino (2.9 km 2 ) and of a 
number of small uninhabited islets (each less than 10 ha). The islands are mainly 
composed of Oligo-Miocene limestones, the soils are young and very similar to the 
parent rocks, and there are no mountains, streams or lakes, but only minor springs. The 
climate is typically Mediterranean and strongly bi-seasonal: the average annual rainfall 
is c. 530 mm, of which some 85% falls during the period October to March; the mean 
monthly temperature range is 12-26°C, and the islands are very windy and sunny. The 
main geomorphological features are karstic limestone plateaux, hillsides covered with 
clay taluses, gently rolling limestone plains, valleys (widien, see below) that drain 
runoff during the wet season, steep sea-cliffs on the south-western coasts, and gently 
sloping rocky shores to the Northeast. The islands have been more or less continuously 
inhabited since 7000 BP and human impact is significant. Presently some 38% of the 
land area is cultivated, c. 25% is built up, and the rest is countryside. 

The terrestrial habitats of the Maltese Islands are mainly characterized by the 
vegetation which can be grouped in three categories: (i) communities that are part of 
the successional sequence (steppe, garigue, maquis) towards a climax (sclerophyll 
forest); (ii) communities which are either specialised to occupy particular habitats, or 
occupy habitats that are rare on the islands, or are relics from a previous ecological 
regime, now surviving in a few réfugia; and (iii) vegetational assemblages of disturbed 
habitats, occupying land subject to periodic disturbance, usually related to anthropic 
activities. 

It is thought that before humans colonised the Maltese Islands, large areas were 
covered with a Mediterranean sclerophyll forest characterised by Quercus ilex and 
Pinus halepensis. The early settlers cut the trees for wood and to clear the land for 
agriculture and buildings, and introduced sheep and goats whose grazing and browsing 
prevents the trees from regenerating. The native forest on the Maltese Islands is all but 
extinct and only remnants persist at four localities, none of which has more than a few 
dozen trees. More extensive tree-covered areas nonetheless exist on the islands; 
however, all owe their origin to human activities (e.g., gardens, plantations, orchards 
etc.). Although originally planted, some are now self-maintaining and self -rege- 
nerating, and therefore qualify as semi-natural woodlands. 

The Maltese maquis is an impoverished scrub community resulting from 
degeneration of the climax woodland due to cutting, grazing and erosion of the soil. A 



CENTIPEDES OF THE MALTESE ARCHIPELAGO 



435 



Gozo 




Cornino 



Malta 



N 

▲ 

10 km 




Filfla 

Fig. 1 
Map of the Maltese Islands showing the localities from where centipedes recorded in this paper 
were collected. Key (in alphabetical order): Malta: 1 Attard, 2 Bahrija (Wied tal-Bahrija), 3 
Ballut ta' 1-Imgiebah, 4 Ballut tal-Wardija/Wardija, 5 Bingemma Gap, 6 Birkirkara, 7 Borg in- 
Nadur (Birzebbuga), 8 Buskett, 9 Chadwick Lakes, 10 Fomm ir-Rih, 11 Ghadira, 12 Ghajn 
Hadid (Selmun), 13 Ghajn Rihana, 14 Ghajn Tuffieha, 15 Ghar Lapsi, 16 Girgenti, 17 Gnejna, 
18 Golden Bay, 19 Gwardamangia, 20 Il-Fawwara (Dingli Cliffs), 21 Il-Maqluba (Qrendi), 22 
Manoel Island, 23 Mgiebah Bay, 24 Mistra Bay/Tal-Kortin, 25 Rabat (St. Agatha's Catacombs), 
26 Salina, 27 San Anton Gardens, 28 San Pawl tat-Targa, 29 Siggiewi, 30 Sliema, 31 Ta' 
Hammud (Mtahleb), 32 Tal-Qroqq (UOM), 33 Targa Gap, 34 Wied Anglu, 35 Wied Babu, 36 
Wied Bufula, 37 Wied Hassabtan, 38 Wied il-Faham, 39 Wied il-Ghasel (Mosta), 40 Wied Incita, 
41 Wied is-Sewda, 42 Wied Qannotta, 43 Wied Znuber, 44 Xrobb 1-Ghagin; Cornino: 45 Central 
area, 46 Santa Marija Bay; Gozo: 47 Dwejra, 48 Fort Chambray slopes, 49 Gharb , 50 Hondoq 
ir-Rummien, 51 Kercem, 52 Qala, 53 Ramla, 54 Ta' Cene, 55 Wied ic-Cawla, 56 Wied il-Mielah. 
57 Xatt 1-Ahmar, 58 Xlendi. 

semi-natural maquis survives in relatively inaccessible sites, such as the sides of steep 
valleys, and at the foot of escarpments, while an artificial maquis develops round 
previously cultivated trees, mainly Olea europaea and Ceratonia siliqua. 



436 M. ZAPPAROLI ETAL. 

The most widespread natural vegetation type present is the garigue. Some 
garigue communities are natural, others result from degradation of forest and maquis, 
particularly where removal of the original vegetation cover has caused such extensive 
soil erosion that large tracts of the limestone bedrock have become exposed and only 
patches of stony soil still occur. Gangues are typical of such rocky ground and are 
especially common on the flat karstic limestone platforms of western Malta and the 
Gozitan hills. Many subtypes of Maltese garigue exist; the principal ones are those 
dominated by Coridothymus capitatus, Anthyllis hermanniae, Teucrium fructicans, 
Erica multiflora, and the endemic Euphorbia melitensis; mixed garigues dominated by 
two, three or more of these species are also common. 

Steppic assemblages dominated by grasses, umbellifers, thistles and geophytes 
are widespread and result from degradation of the maquis and garigue, due to grazing 
and browsing and from soil erosion due to the short but heavy rainstorms which are 
characteristic of the islands. Some steppic communities are, however, climactic or 
semi-climactic with Lygeum spartum on clay slopes, or with Hyparrhenia hirta and 
Andropogon distachyus. Other steppes are characterised by Brachypodium retusum or, 
rarely, by Phalaris truncata. The more degraded steppes are characterised by Stipa 
capensis and Aegilops geniculata and a variety of thistles (e.g., Carlina involucrata, 
Notobasis syriaca, Galactites tomentosa) and geophytes (e.g., Asphodelus aestivus, 
and Urginea pancration). Steppic communities also develop on abandoned agricultural 
land, which is increasing in extent. 

Habitats that are not part of the successional sequence include coastal marsh- 
lands, sand dunes, maritime vegetation, freshwater, and rupestral communities. 

Maltese coastal marshes are characterized by a muddy substratum on which a 
pool of brackish water collects in the wet season. During the dry season this water 
becomes progressively more brackish until it finally disappears completely, leaving the 
marsh dry until the following wet season. 

Many local sandy beaches were backed by dune systems, but at present only 
very few persist and even these have been much degraded mainly due to human 
activities in connection with beach development for touristic purposes and recreational 
use. Sand dune ecosystems are thus amongst the rarest and most threatened of local 
ecosystems. Local dunes are dominated by the dune grasses Elytrigia juncea and 
Sporobolus pungens, and, until recently, also by Ammophila littoralis which has now 
been totally extirpated. 

On gently sloping rocky shores halophytic vegetation grows in isolated patches 
on the shallow saline soil that accumulates in pockets of the rock. The species present 
form part of the Mediterranean vegetational community called the Crithmo- 
Limonietum. 

Rupestral assemblages dominated by shrubs occur on sheer rock faces and 
cliff/scree environments, mainly at the south, southwest and west coasts of the islands. 
Because of their relative inaccessibility these habitats provide important refuges for 
many species of Maltese flora and fauna, including many endemics, amongst which are 
two plant taxa (Palaeocyanus crassifolius and Cremnophyton lanfrancoi) belonging to 
monotypic genera. 

The main freshwater habitats are those associated with valleys (in Maltese: 
widien), which are géomorphologie ally dry valleys, that is, valleys formed during a 



CENTIPEDES OF THE MALTESE ARCHIPELAGO 437 

previous climatic regime (the Pleistocene pluvial periods), which are now dry for some 
months of the year and in which water only flows during the wet season. However, 
some local widien drain springs originating from perched aquifers and retain some 
surface water even during the dry season. Other freshwater habitats are temporary 
rainwater pools, formed by rainwater that collects in natural depressions and hollows 
in rock during the wet season, and a few permanent ponds. 

In spite of being made up almost exclusively of limestone, the Maltese Islands 
have surprisingly few known deep caves. Those caves that have been explored bio- 
logically have revealed an impoverished but interesting biota with a number of 
endemic invertebrates. 

Because of the islands' high human population and considerable land use, 
anthropogenic habitats have a large coverage. Such habitats are dominated by a variety 
of plant species consisting mainly of ruderals and aliens. Different types occur in 
association with agriculture, afforestation, abandoned fields, along roadsides, in 
disturbed seaside habitats and in urban areas. 

The above synthesis is based on Alexander (1988), Axiak et al. (2002), Bowen 
Jones et al. (1961), Chetcuti et al. (1992), Haslam et al. (1977), Lanfranco (1995), 
Pedley et al. (1976), Schembri (1993, 1997), Schembri et al. (1999), Schembri & 
Lanfranco (1993), and Vossmerbäumer (1972); these works should be consulted for 
more detail and for an extensive bibliography. 



MATERIAL AND METHODS 

The present paper is based on literature records that have been critically revised, 
and on unpublished material. For each species the following is reported: the scientific 
name; the complete name of the author and year of publication; the bibliographic 
references concerning the study area listed chronologically, with the name of the 
species and author as originally quoted; the general geographic distribution, mainly as 
a list of the countries or geopolitical units from where the species is known, critically 
revised from the literature; the chorotype according to Vigna Taglianti et al. (1992, 
1999); the list of collecting sites for the species in the Maltese Islands, arranged 
according to island (Malta, Cornino, Gozo), with the collecting sites for each island 
listed alphabetically; a synthesis of available data on habitat preferences in the study 
area; and taxonomic remarks where relevant. 

The following acronyms are used for the collectors of the material examined 
and for the collections where this material is now deposited. Collectors: AD = A. 
Deidun; AV = A. Valle; CA = Causin; DC = D. Caruso; DJ = D.M. Johnson; EG = E.H. 
Giglioli; EL = E. Lanfranco; GT = G.B. Toscanelli; JS = J.L. Schembri; LM = L. Main; 
MG = M. Gauci; MP = M. Pace; PS = P.J. Schembri; SA = S. Azzopardi; SS = S. 
Schembri; ST = S. Saliba. Collections: AM = A. Minelli; BG = Museo Civico di 
Scienze naturali "E. Caffi", Bergamo, Italy; CT = Dipartimento di Zoologia, Università 
di Catania, Italy; DBUM = Department of Biology, University of Malta; FI = Museo 
di Storia naturale di Firenze, sezione Zoologica "La Specola"; MHNG = Muséum 
d'histoire naturelle, Genève; MZ = M. Zapparoli. 



438 M. ZAPPAROLI ETAL. 



LIST OF THE SPECIES 

Scutigeromorpha Gervais, 1837 
Scutigeridae Gervais, 1837 

Scutìgera Lamarck, 1801 

1. Scutìgera coleoptrata (Linné, 1758) 

Cermatia variegata Risso: Gulia, 1890: 41. 
Cermatia variegata: Gulia, 1913: 554. 
Scutigera coleoptrata: Schembri, 1996: 120. 

Material examined. Malta: 1, Ballut ta' 1-Imgiebah, 8.4.1984, PS (MZ, MZ det.); 1, 
Chadwick Lakes, 7.3. 1975, DC (CT, AM det.); l,G'Mangia, 1987,MG(MZ,MZdet.); 1, Malta, 
5.1973, AV (BG, AM det.); 1, Manoel Island, 16.3.1985, PS (MZ, MZ det.); 1, S. Antonino 
[= San Anton Gardens], 12.3.1975, leg. ? (CT, AM det.); Siggiewi, 6.1982, MP (MZ, MZ det). 

General distribution. Portugal, Spain (incl. Balearic Is.), France (incl. Corsica), 
Italy (incl. Sardinia and Sicily), Switzerland, S. Germany, Austria, Czech Republic, 
Slovakia, S. Hungary, Slovenia, Croatia, Montenegro, FYR Macedonia, Albania, main- 
land and insular Greece (incl. Crete), Bulgaria, Romania, Ukraine (incl. Crimea), 
Caucasus, Near and Middle East, N. Africa (Egypt, Libya, Tunisia, Algeria, Morocco); 
also recorded from Madeira and the Canary Is. (introduced ?); introduced in Central 
and N. Europe, Asia, N. America, S. Africa, St. Helena, Argentina (Attems, 1907; 
Würmli, 1973, 1977). 

Chorotype. Mediterranean (MED). 

Ecological notes. Widespread species on the Maltese Islands, occurring in a 
range of habitats, usually with some degree of humidity, including human habitations 
and their surroundings (leaf litter, under thick vegetation, under stones on soil, in 
cellars etc). 

Lithobiomorpha Pocock, 1895 
Lithobiidae Newport, 1 844 

Eupolybothrus Verhoeff, 1907 
Subgenus Allopolybothrus Verhoeff, 1907 

2. Eupolybothrus (Allopolybothrus) nudicornis (Gervais, 1837) 
Eupolybothrus elongatus (Newp.): Matic et ah, 1967: 197. 

Material examined. Malta: 7, Attard, 5.3.1975 (CT, AM det.); 1 o\ Bahrija Valley, 

25.3.1984, PS & SS (DB UM, MZ det.)l 9 , Bingemma Gap, 24.3.1984, SS (MHNG, MZ det.); 
1, Birkirkara, 10.1969, CA (MZ, MZ det.); 1, Buskett, 12.1.1980, PS (MZ, MZ det.); 1, 
Chadwick Lakes, 23.4.1983, PS (MZ, MZ det.); 1, Ghadira, 27.12.1979, PS (MZ, MZ det.); 1 
juv., Ghajn Hadid, Selmun, 5.12.1983, MG (MHNG, MZ det.); 1, Ghar Lapsi, 6.3.1975, DC (CT, 
AM det.); 2, Malta, 2.3. 1975, leg. ? (CT, AM det.); 4 S , 1 2,1 9 juv., Manoel Island, 16.3. 1985, 
PS (MZ, MZ det.); 1, Mistra Bay, 3.3.1975, DC (CT, AM det.); 1 8, Rabat, St. Agatha's 
Catacombs. 16.7.1984, MG (MHNG, MZ det.); 2 S, San Pawl tat-Targa, garden, pitfall trap, 

17.2.1985, DJ (DBUM, MZ det.); 1 9, Tal-Kortin, Mistra, 18.3.1984, PS (MZ, MZ det.); 2 o\ 
1 9 , Tal-Qroqq, UOM, under stones, Acacia stand, 27.3.1984, SA & LM (MZ, MZ det.); 4 o\ 
1 9 , ibidem, pitfall trap, Acacia stand, 5.4.1984, SA & LM (MHNG, MZ det.); 5 6,2 9 , ibi- 
dem, carob. under stones, 16.4.1984, SA & LM (MZ, MZ det.); exx., Targa Gap, 10.1.1970, 



CENTIPEDES OF THE MALTESE ARCHIPELAGO 439 



leg.?; 3 juv., Wied Anglu, under stones, in watercourse, 16.12.1984, PS (MHNG, MZ det.); 2, 
Wied Babu, 23.5.1985, SS (MZ, MZ det.); exx., Wied Hassabtan, 20.1.1970, leg.? (MZ, MZ 
det.); 2 o\ 1 5, Wied Incita, 25.3.1984, PS & SS (DBUM, MZ det.); 1, Wied is-Sewda, 
26.3.1975, SS (MZ, MZ det); 1 3 juv., 2 9 juv., Wied il-Ghasel, Mosta, garigue, under stones, 
26.1.1985, PS (DBUM, MZ det.). Gozo: 1, Dwejra, 7.3.1975, DC (CT, AM det.); 1 9 juv., 1 
imm., limits of Gharb, 14.2.1985, PS (MHNG, MZ det.); l,Qala, 13.3.1975, DC (CT, AM det.); 
2 9 juv., Wied il-Mielah,16.2.1985, PS (DBUM, MZ det.). 

General distribution. SE. France (Basses Alpes, Alpes Maritimes), Corsica, 
Italy (Apennines), Sardinia, Sicily, Malta, NE. Morocco, N. Tunisa, N. Algeria 
(Brolemann, 1921, 1930; Foddai et al, 1995; Matic et al, 1967; Zapparoli, 1984). 
Reported also from Spain by Attems (1927, 1952), but record needs to be confirmed. 

C horotype. W-Mediterranean (WME). 

Remarks. The old records of Gulia (1890, 1913) quoted under Lithobius forfi- 
catus (Linnaeus, 1758) may refer to this species (see below). 

Ecological notes. Common and widespread species on the Maltese Islands, 
occurring in a range of habitats that include widien, leaf litter under trees such as 
Acacia and Ceratonia siliqua, under stones in garigue, coastal vegetation, gardens and 
urbanised areas. 

Lithobius Leach, 1814 
Subgenus Lithobius Leach, 1814 

3. Lithobius (Lithobius) castaneus Newport, 1844 

Material examined. Malta: 1 9, Ghajn Rihana, 25.3.1984, PS (MHNG, MZ det.); 1, 
Malta, 2.3.1975, DC (CT, AM det.); 1, Malta, 5.4.1978, leg.? (MZ, MZ det.); 1, Il-Maqluba, 
20.1.1980, SS (MZ, MZ det.); 1, ibidem, 14.2.1982, PS (MZ, MZ det.); 1 S juv., San Pawl tat- 
Targa, garden, pitfall trap, 3.2.1985, DJ (MHNG, MZ det.); 1 o\ 1 9 , Tal-Qroqq, UOM, under 
stones, Acacia stand, 27.3.1984, SA & LM (DBUM, MZ det.); 3, ibidem, pitfall trap, Acacia 
stand, 5.4.1984, SA & LM (MZ, MZ det.); 6 9 , ibidem, carob, under stones, 16.4.1984, SA & 
LM (MZ, MZ det.); 1 9, Wied Babu, 23.5.1985, SS (MHNG, MZ det.); 1 9, Wied 1-Ghasel, 
Mosta, garigue, under stones, 26.1.1985, PS (DBUM, MZ det.). 

General distribution. Morocco, Tunisia, Algeria, Portugal, Spain, France (incl. 
Corsica), Italy (including Sardinia and Sicily), Malta, S. Austria, Slovenia, Croatia, 
Bosnia Herzegovina, Serbia (Brolemann, 1921, 1930, 1932;Eason, 1982; Foddai et al, 
1995; Kos, 1992; Machado, 1952; Matic et al, 1967; Stoev, 1997; Zapparoli, 1981); 
the species has been reported from Bulgaria, but records require confirmation (Stoev, 
2002); introduced in Guatemala (Eason, 1973). 

Chorotype. S-European (SEU). 

Ecological notes. Common and widespread; occurring in a variety of habitats 
including widien, leaf litter under trees, under stones in garigue, and in gardens. 

4. Lithobius (Lithobius) forficatus (Linnaeus, 1758) 

Lithobius forficatus Leach [sic]: Gulia, 1890: 41. 
Lithobius forficatus: Gulia, 1913: 554. 

Material examined. No material from Malta examined (cf. Remarks). 
General distribution. Iceland (introduced), Finland, Norway, Sweden, Den- 
mark, United Kingdom, Ireland, Netherlands, France (incl. Corsica, but introduced), 



440 M. ZAPPAROLI ETAL. 

Germany, Poland, Czech Republic, Slovakia, Hungary, Switzerland, Austria, Italy 
(incl. Sardinia and Sicily (Eolian Is.), but introduced in both), Slovenia, Croatia, 
Bosnia Herzegovina, Serbia, Montenegro, FYR Macedonia, Romania, Bulgaria, 
Albania, mainland Greece, N. Turkey, Georgia, Russia (Krasnodar Prov.); also present 
in N. Africa (introduced?) and in Malta (introduced); from the W-Palearctic it has been 
introduced to N. America, S. America, St. Helena, Hawaii Is. (established?), Kuriles 
(Bròlemann, 1930; Eason, 1964, 1970, 1982, 1996; Enghoff, 1983; Foddai et al, 1995; 
Kos, 1992; Orszâgh, 2001; Stoev, 1997; Tajovsky, 2001; Wytwer, 1997; Zalesskaja, 
1978; Zapparoli, 1999). 

Chorotype. European (EUR). 

Remarks. According to Zapparoli (1995a) this species has a very limited distri- 
bution in the micro- and macro-insular systems of the Mediterranean area, although it 
is highly anthropophilous. L. forficatus has never been recorded from the Balearic Is., 
Corsica, Sardinia, Sicily, Crete and Cyprus, from where it must be considered absent. 
In the Tyrrhenian area only a few records from Capri, the Eolian Is. (Lipari, Vulcano) 
and Lampedusa are known, all probably resulting from anthropic introductions. The 
only record of this species from the Maltese Islands is that of Gulia (1890, 1913), 
however, in spite of the in-depth research carried out no new records have been added 
since the end of the 19th century and we suspect that Gulia's record from Malta may 
be based on a misidentification and probably refers to another lithobiid species 
(possibly E. nudicornis ?). 

Ecological notes. No data available. 

5. Lithobius (Lithobius) lapidicola Meinert, 1872 

Material examined. Malta: 1 6, Buskett, leaf litter on tree, 30.12.1978, SS (MZ, MZ 
det.); 4 o\ 3 ?, Sliema, 22/23.12.1969, EL (MZ, MZ det.). 

General distribution. Canary Is., Ireland, United Kingdom, Sweden, Nether- 
lands, Germany, Switzerland, Denmark, Poland, Czech Republic, Slovakia, Ukraine, 
France (incl. Corsica), Italy (incl. Sardinia and Sicily), Austria, Hungary, Slovenia, 
Bosnia Herzegovina, Montenegro, Romania, Albania, mainland Greece (incl. Ionian 
Is.) (Bròlemann, 1930; Eason, 1964, 1970, 1982, 1985, 1996; Enghoff, 1983; Foddai 
et al, 1995, 1996; Kos, 1992; Orszâgh, 2001; Stoev, 1997; Tajovsky, 2001; Wytwer, 
1997; Zalesskaja, 1978). 

Chorotype. European (EUR). 

Ecological notes. Records available at present indicate that this is a leaf litter 
species which occurs in semi-natural wooded areas (Buskett) and in urban gardens 
(Sliema). 

6. Lithobius (Lithobius) peregrinus Latzel, 1880 

Material examined. Malta: 1 o\ Malta, date?, SS (MZ, MZ det.). 

General distribution. SE. Italy (Gargano), Bosnia Herzegovina, Montenegro, 
FYR Macedonia, Albania, mainland Greece (incl. Ionian Is.), Bulgaria, Caucasus; 
introduced in the United Kingdom, France, Spain, NE. Italy, S. Africa, Bermuda Is., 
Panama (Stoev, 1997, 2001; Zapparoli, 1992). 



CENTIPEDES OF THE MALTESE ARCHIPELAGO 441 

Chorotype. S-European (SEU). 

Remarks. There are no previous records of this species from Malta, were it has 
probably been introduced. 

Ecological notes. No data available. 

7. Lithobius (Lithobius) trinacrius Verhoeff, 1925 

Material examined. Malta: 1 <?, Ballut ta' 1-Imgiebah, 8.4.1984, PS (MHNG, MZ det.); 
1 6, Ghajn Hadid, 8.5.1983, PS (MZ, MZ det.); 1 9, Ghajn Hadid, Selmun, 5.12.1983, MG 
(MHNG, MZ det.); 19,2 juv., Selmun, 5.2.1983, PS (MZ, MZ det.); 2 3,2 9, Mgiebah, 
9.11.1985, leg. ? (MZ, MZ det.). Gozo: 1 o\ Dwejra, 9.2.1984, MG (MHNG, MZ det.); 1 o\ 1 
9 , Ramla, 15.2.1986, clay slopes, PS (DBUM, MZ det.); 1 6, Xlendi, 26.4.1984, MG (MZ, MZ 
det.). 

General distribution. Sicily (Foddai et al., 1995); also reported from Pantelleria 
(Zapparoli, 1995a). 

Chorotype. W-Mediterranean (WME). 

Ecological notes. Available records indicate that this species has a distribution 
limited to coastal areas where it occurs under trees, shrubs, in grass steppes, on clay 
slopes and under maritime vegetation. 

Subgenus Monotarsobius Verhoeff, 1905 

8. Lithobius (Monotarsobius) crassipes L. Koch, 1862 

Material examined. Malta: 2 â , Ballut tal-Wardija, soil and leaf litter, Berlese extractor, 

8.4.1984, PS & SS (MZ, MZ det.); 1 «J, 1 9, Nadur, near Bingemma Gap, 30.12.1984, PS 
(MHNG, MZ det.); 1 9, Wied Qannotta, S.E. end, 11.12.1983, PS (DBUM, MZ det.); 1 <J juv., 
Wied 1-Ghasel, Mosta, 26.1.1985, PS (DBUM, MZ det.); 1 ö\ Wied il-Faham, 27.2.1982, PS 
(MZ, MZ det.). Gozo: 2 ó\ 5 9 , limits of Kercem, 14.2.1985, PS (MZ, MZ det.); 1 ò\Ta' Cene, 

2.2.1985, PS & MG (MHNG, MZ det.); 1 o\ 1 9 juv., Wied ic-Cawla, 9.4.1984, MG (MZ, MZ 
det.); 3 S, 1 9, Wied il-Mielah, 16.2.1985, PS (MZ, MZ det.). 

General distribution. Scandinavia, United Kingdom, Ireland, Iberia, France, 
Netherlands, Germany, Switzerland, Poland, Czech Republic, Slovakia, Austria, Italy 
(incl. Sardinia and Sicily), Slovenia, Croatia, Bosnia Herzegovina, Montenegro, 
Serbia, Albania, FYR Macedonia, mainland and insular Greece (incl. Crete), Bulgaria, 
Romania, Russia, Turkey, Syria, Jordan, Central Asia, Algeria, Tunisia, Canary Is., 
Madeira; probably introduced in N. America (Brölemann, 1921, 1930, 1932; Eason, 
1964, 1982, 1985; Foddai et al, 1995; Kos, 1992; Marie et al, 1967; Orszâgh, 2001; 
Stoev, 1997, 2001; Tajovsky, 2001; Wytwer, 1997; Zalesskaja, 1978; Zapparoli, 1991, 
1999). 

Chorotype. W-Palaearctic (WPA). 

Ecological notes. Common and quite widespread; this species occurs in soil and 
leaf litter beneath trees and shrubs, especially in sheltered situations, such as maquis in 
the deeper widien. 

SCOLOPENDROMORPHA PoCOCk, 1895 

Scolopendre} AE Newport, 1 844 
Scolopendra Linné, 1758 



442 M. ZAPPAROLI ETAL. 

9. Scolopendra cingulata Latreille, 1829 

Scolopendra cingulata Latr.: Gulia, 1890: 41. 
Scolopendra cingulata: Gulia, 1913: 554. 
Scolopendra cingulata Latr.: Matic et al., 1967: 197. 
Scolopendra cingulata: Schembri, 1996: 120. 

Material examined. Malta: 1, Wied Mejxu, under stones, 25.10.1970, EL (MZ det.); 1, 
Birkirkara, under stones, 5.11.1974, SS (MZ det.). Cornino: 2, under stones, 23.03.1975, SS (MZ 
det.); 1, Santa Marija Bay, 26.9.1982, PS (MZ, MZ det.). 

General distribution. Tunisia, Algeria, Morocco, Portugal, Spain, France, Italy, 
Slovenia, Croatia, Bosnia Herzegovina, Serbia, Montenegro, FYR Macedonia, 
Romania, Bulgaria, Albania, mainland and insular Greece, Turkey, Hungary, Ukraine, 
S. European Russia (Crimea, Caucasus), NW Iran, Syria, Lebanon, Palestine, Israel, 
Jordan, Egypt (Sinai), Cyrenaica; also in Sicily and Cyprus; absent in Balearic Is., 
Corsica, Sardinia and Crete (Attems, 1930; Bròlemann, 1921, 1930, 1932; Foddai et al, 
1995; Kos, 1992; Lewis, 1985; Serra, 1983; Stoev, 1997; Zalesskaja & Schileyko, 1992; 
Zapparoli, 1991, 1999). Two records from Tadjikistan (Zalesskaja & Schileyko, 1992). 

Chorotype. (Turano?)-Mediterranean (MED). 

Remarks. Recorded from many localities on Malta, as well as from the islands 
of Cornino and Gozo (Matic et al., 1967); many more specimens were seen by one of 
us (PS) but were not collected. 

Ecological notes. Widespread species on the Maltese Islands, occurring in a 
wide range of habitats including coastal garigue and clay slopes, widien, inland garigue 
and fields. 

10. Scolopendra oraniensis Lucas, 1846 

Scolopendra canidens oraniensis (Luc.) [sic]: Matic et al., 1967: 197. 

Material examined. Malta: 1, Attard, 5.3.1975, leg. ? (CT, AM det.); 1, Bahrija Valley, 
6.4.1985, PS (MZ, MZ det.); 1, Ghadira, 4.3.1975, leg. ? (CT, AM det.); 1, Ghajn Tuffieha, clay 
slopes, 24.2.1985, SS (MZ, MZ det.); 1, Ghar Lapsi, 6.3.1975, DC (CT, AM det); 1, Gnejna, 
104.1982, JS (DBUM, MZ det.); 1, Il-Fawwara, Dingli cliffs, 9.4.1983, SP (MZ, MZ det.); 1, 
Salina, 5.5.1984, SS (MZ, MZ det); 1, Mgiebah Bay, clay slopes, 8.4.1984, PS (MZ, MZ det); 
1, Mistra, 3.3.1975, DC (CT, AM det); 1, Wardija, 2.3.1975, leg. ? (CT, AM det.); 1, Wied 
Bufula, 8.11.1986, leg. ? (MZ, MZ det); 1, Wied Incita, 25.3.1984, PS & SS (DBUM, MZ det); 
1, Wied 1-Ghasel, Mosta, garigue, under stones, 26.1.1985, PS (MZ, MZ det); 1, Wied Znuber, 
15.4.1984, PS (MHNG, MZ det.); 1, Xrobb 1-Ghagin, 7.5.1984, MG (MHNG, MZ det). Gozo: 
1, Dwejra, 17.5.1984, SS (MHNG, MZ det.); 2, Hondoq ir-Rummien, 5.3.1984, SS (DBUM, MZ 
det.); 1, Wied il-Mielah, 9.4.1984, MG (MZ, MZ det). 

General distribution. Portugal, Spain (incl. Balearic Is.), S. France, Corsica, 
Central and S. Italy, Sardinia, Sicily, Malta. Records from Morocco and Algeria are 
also known (Wiirmli, 1980). 

Chorotype. Western Mediterranean (WME). 

Ecological notes. Common and widespread especially in coastal areas; it occurs 
in widien, garigue and on clay slopes. 

Cryptopidae Kohlrausch, 1881 
Cryptops Leach, 1815 

11. Cryptops trisulcatus Bròlemann, 1902 
Cryptops trisulcatus Brol. [sic]: Matic et al., 1967: 197. 



CENTIPEDES OF THE MALTESE ARCHIPELAGO 443 



Material examined. Malta: 1, Chadwick Lakes, 7.3.1975, DC (CT, AM det.); 1, Ghadira, 
4.3.1975, DC (CT, AM det.). Gozo: 1, Dwejra, 9.11.1984, MG (MZ, MZ det.); 1, ibidem, 
17.5.1984, SS (MZ, MZ det.). Cornino: 1, central area, 26.9.1982, PS (MZ, MZ det.). 

General distribution. Canary Is., Algeria, Portugal, Spain (incl. Balearic Is.), S. 
France (incl. Corsica), Italy (Apenninic), Sardinia, Sicily, Romania, insular Greece 
(Ionian Is., S. Sporades, Crete), SE. Turkey (Attems, 1930; Brölemann, 1921, 1930, 
1932; Foddai et al, 1995, 1996; Machado, 1952; Negrea & Matic, 1973; Stoev, 1997; 
Zapparoli, 1999). 

Chorotype. Mediterranean (MED). 

Ecological notes. Available records suggest that this species has a rather limit- 
ed distribution and occurs mostly in coastal areas, but also in widien (Chadwick 
Lakes). 

Geophilomorpha Leach, 1815 

HlMANTARIIDAE Cook, 1895 

Himantarium C. Koch, 1847 

12. Himantarium gabrielis (Linnaeus, 1767) 

Himantarium gabrielis: Schembri, 1996: 120. 

Material examined. Malta: 1, Ballut ta' 1-Imgiebah, 8.4.1984, SS (MZ, MZ det.); 1, 
Ballut tal-Wardija, 8.4.1984, PS (MHNG, MZ det.); 1, Borg in-Nadur, B 'Bugia, 20.1.1985, PS 
(MZ, MZ det.); 1, Il-Maqluba, Qrendi, 7.12.1982, PS (MHNG, MZ det.); 3, Malta, 
12.1972/1.1973, AV (BG, AM det.); 1, San Pawl tat-Targa, 3.1983, DJ (MZ, MZ det.); 1, San 
Pawl tat-Targa, garden, pitfall trap, 17.2.1985, DJ (MHNG, MZ det.); 1, Tà Hammud, Mtahleb, 
18.2.1983, PS (MZ, MZ det.); 1, Tal-Qroqq, UOM, 16.4.1984, SA & LM (MHNG, MZ det.); 1, 
Wied Anglu, 16.12.1984, PS (DBUM, MZ det.); 1, Wied Bufula, 8.11.1986, leg. ? (MZ, MZ 
det.). 

General distribution. Tunisia, Algeria, Morocco, S. France (incl. Corsica), Italy 
(incl. Sardinia and Sicily), Slovenia, Croatia, Bosnia Herzegovina, Montenegro, FYR 
Macedonia, Albania, mainland and insular Greece (excl. Crete), S. Romania, Bulgaria, 
W. Turkey; introduced in Madagascar (Attems, 1929; Brölemann, 1921, 1930, 1932; 
Foddai et a/., 1995; Kos, 1992; Minelli et al, 1984; Stoev, 1997, 2001; Zapparoli, 
1999). Reported from Portugal by Attems (1929) but not by Machado (1952); also 
occurring in Central Europe (Brölemann, 1930). 

Chorotype. Mediterranean (MED). 

Ecological notes. Common and widespread species on the Maltese Islands, 
occurring in a wide range of habitats that include leaf litter under trees and shrubs, soil 
in garigue, widien and gardens. 

Stigmatogaster Latzel, 1880 

13. Stigmatogaster gracilis (Meinert, 1870) 

Material examined. Malta: 2, 12.1972/1.1973, AV (BG, AM det.); 2, Malta, 8.1878, EG 
(FI, AM det.). 

General distribution. Tunisia, Algeria, Balearic Is., S. France (incl. Corsica), 
Italy (incl. Sardinia and Sicily), Croatia, Montenegro, Albania, mainland and insular 



444 M. ZAPPAROLI ET AL. 

Greece (excl. Crete) (Brölemann, 1921, 1930, 1932; Foddai et al, 1995; Kos, 1992; 
Negrea & Matic, 1973; Stoev, 1997). 

Chorotype. Mediterranean (MED). 

Ecological notes. No data available. 

Bothriogaster Sseliwanoff, 1879 

14. Bothriogaster signata (Kessler, 1874) 
Bothriogaster signata Att. [sic]: Matic et al., 1967: 196. 

Material examined. Malta: 3, Tal-Qroqq, UOM, Acacia stand, under stones, 27.3.1984, 
SA & LM (MZ, MZ det.). Gozo: 2, Dwejra, 17.5.1984, SS (DBUM, MZ det.); 1, Qala, 
17.5.1985, MG (MHNG, MZ det.). 

General distribution. FYR Macedonia, Albania, Bulgaria, mainland and insular 
Greece (incl. Crete), Turkey, Cyprus, Syria, Palestine, Israel, Egypt, Libya (Cyrenaica, 
Tripolitania), Tunisia, Caucasus, Iran, Iraq, Jordan, Saudi Arabia, Turkestan, 
Usbekistan (Stoev, 2000; Zapparoli, 1991). 

Chorotype. Turano-Mediterranean (TUM). 

Remarks. Previously recorded from several localities on the island of Malta, 
as well as from Cornino and Gozo (Matic et al., 1967). 

Ecological notes. Only few records are available but the species seems to pre- 
fer soil under vegetation in rather arid situations. 

DlGNATHODONTIDAE Cook, 1895 

Dignathodon Meinert, 1870 

15. Dignathodon microcephalus (Lucas, 1846) 
Dignathodon microcephalum [sic] (Luc): Matic et al., 1967: 197. 

Material examined. Malta: 2, Targa Gap, 2.10.1982, PS & SS (MZ, MZ det.). 

General distribution. Morocco, Algeria, Tunisia, Portugal, Spain (incl. Balearic 
Is.), S. France (incl. Corsica), Italy (incl. Sardinia and Sicily), Austria, Croatia, Bosnia 
Herzegovina, Serbia, Montenegro, Czech Republic, Slovakia, Romania, Bulgaria, 
Albania, mainland and insular Greece (incl. Crete), Near and Middle East, Crimea; al- 
so recorded from Luxembourg, where it was probably introduced (Attems, 1929 
Brölemann, 1921, 1930, 1932; Dobroruka, 1956; Foddai et al., 1995, 1996; Kos, 1992 
Machado, 1952; Matic, 1972; Negrea & Matic, 1973; Orszâgh, 2001; Stoev, 1997 
Tajovsky, 2001; Zapparoli, 1991, 1995b, 1999). 

Chorotype. Mediterranean (MED). 

Remarks. This species has been previously recorded from Gozo (Matic et al., 
1967). 

Ecological notes. No data available. 

Henia C.L. Koch, 1847 

Subgenus Meinertia Bollmann, 1893 

16. Henia (Meinertia) bicarinata (Meinert, 1870) 



CENTIPEDES OF THE MALTESE ARCHIPELAGO 445 

Material examined. Malta: 1, Bahrija Valley, 25.3.1984, PS & SS (MHNG, MZ det.); 1, 
Buskett, soil, leaf litter, Berlese extractor, 30.4.1984, PS (MZ, MZ det.); 1, Chadwick Lakes, 
7.3.1975, DC (AM, AM det.); 1, Fomm ir-Rih (cobble beach), October 2001, MG (AM, AM 
det.); 1, Fomm ir-Rih (cobble beach, on wrack), October 2001, MG (AM, AM det.); 1, Wied 1- 
Ghasel, near Mosta Fort, 16.3.1985, PS (MHNG, MZ det.); 1, Manoel Island, 3.4.1984, MG 
(MZ, MZ det.); 2, Mgiebah, 17.3.1985, leg. ? (DBUM, MZ det.). Gozo: 1, Dwejra, 17.5.1984, 
SS (MZ, MZ det.); 1, Fort Chambray slopes, 3.2.1985, PS & MG (MHNG, MZ det.); 1, Ta' 
Cene, 2.2.1985, PS & MG (DBUM, MZ det.); 1, Tà Cene, 11.3.1975, DC (AM, AM det.). 

General distribution. Macaronesia, Maghreb, Iberia, France (incl. Corsica), 
Italy (peninsular regions, Sardinia and Sicily), Croatia, Bosnia Herzegovina, Slovakia, 
Hungary, Bulgaria, mainland and insular Greece (incl. Crete), Turkey, Caucasus 
(Minelli, 1982; Orszâgh, 2001). 

Chorotype. Mediterranean (MED). 

Ecological notes. A more or less widespread species that occurs in coastal areas 
in stranded Posidonia debris on beaches, on coastal clay slopes and amongst coastal 
vegetation, but also inland in widien, in soil and leaf litter under trees and shrubs, and 
under stones in garigue. 

Subgenus Chaetechelyne Meinert, 1870 

17. Henia (Chaetechelyne) vesuviana (Newport, 1845) 

Material examined. Malta: 4, Nadur, near Bingemma Gap, 30.12.1984, PS (MZ, MZ 
det.). 

General distribution. Tunisia (?), Spain (?), S. France, Corsica (?), Switzerland, 
Italy (incl. Sardinia and Sicily), Croatia, Slovenia, SW. Romania (Minelli, 1982; Kos, 
1992; Stoev, 1997). 

Chorotype. Western Mediterranean (WME). 

Ecological notes. The only locality where this species has been collected on the 
Maltese Islands has mainly a garigue and low maquis vegetation. 

Schendylidae Verhoeff, 1908 
Schendyla Bergsoe & Meinert, 1 866 

18. Schendyla sp. n. 

Material examined. Malta: 7, Ballutta' 1-Imgiebah, soil leaf litter, 8.4.1984, PS (MZ, MZ 
det.); 1, Nadur, near Bingemma Gap, 30.12.1984, PS (MZ, MZ det.); 1 9, Mtahleb, 6.3.1975, 
DC (AM, AM det.). 

Remarks. This new taxon will be described in the context of a revision of the 
Mediterranean Schendyla species. 

Ecological notes. The specimens were found in soil and leaf litter under trees 
(Quercus ilex at Ballut ta' 1-Imgiebah) and in low maquis (Ceratonia siliqua at 
Bingemma Gap). 

Geophilidae Cook, 1895 
Pachymerium C.L. Koch, 1847 

19. Pachymerium ferrugineum (C.L. Koch, 1835) 
Pachymerium ferrugineum (C. Koch) [sic]: Matic et al., 1967: 197. 



446 M. ZAPPAROLI ETAL. 



Material examined. Malta: 1, Ghadira, near reserve, 21.4.1984, PS (MHNG, MZ det.); 
1, Girgenti, 18.10.1986, PS (MZ, MZ det.); 1, Malta, 8.10.1878, EG & GT (FI, AM det.); 1, 
Salina, 5.5.1984, SS (DBUM, MZ det.); 1, ibidem, 22.1.1985, MG (MZ, MZ det.); 2, San Pawl 
tat-Targa, garden, pitfall trap, 17.2.1985, DJ (MZ, MZ det.); 4, Tal-Qroqq, UOM, 5.4.1984, SA 
& LM (MZ, MZ det.); 1, Wied Bufula, 8.11.1986, leg. ? (DBUM, MZ det.); 1, Wied 1-Ghasel, 
Mosta, 16.12.1983, PS (MHNG, MZ det.). Gozo: 1, Fort Chambray slopes, 3.2.1985, PS & MG 
(MHNG, MZ det.); 1, Qala, 17.5.1985, MG (MZ, MZ det); 1, Ramla Dunes, 16.5.1984, SS 
(MZ, MZ det.); 1, Ramla, 4.3.1984, SS (MHNG, MZ det); 1, ibidem, 2.2.1985, PS (MHNG, MZ 
det); 2, ibidem, 15.2.1985, PS (MZ, MZ det); 1, Ta' Cene, 2.2.1985, PS & MG (MZ, MZ det.), 
1, Xatt 1-Ahmar, 4.5.2002, AD (AM, AM det.). 

General distribution. Macaronesia (Azores, Madeira, Canary Is.), N. Africa 
(Tunisia; Algeria incl. Mediterranean coasts and Hoggar, Central Sahara; Morocco; 
Libya: Cyrenaica, Tripolitania), Portugal, Spain (incl. Balearic Is.), France (incl. 
Corsica), Italy (incl. Sardinia and Sicily), Austria, former Czechoslovakia, Poland, 
Latvia, Hungary, Slovenia, Croatia, Bosnia Herzegovina, FYR Macedonia, Albania, 
Bulgaria, Greece, Romania, European Russia, Turkey, Cyprus, Palestine, Iran, 
Caucasus, Turkestan; recorded from Scandinavia (Finland, Norway), United Kingdom, 
Netherlands; also present in Alaska and Pribilof Is.; introduced to Japan, Hawaii Is., N. 
America, Juan Fernandez Is., Mexico, Easter I. (Barber, 1985; Eason, 1964; Meidell, 
1977; Palmén & Rantala, 1954; Stoev, 2000). 

Chorotype. W-Palearctic (WPA). 

Ecological notes. Widespread species on the Maltese Islands, occurring in a 
wide range of habitats including coastal garigue and clay slopes, sandy beaches, 
widien, inland garigue and gardens. 

Clinopodes C.L. Koch, 1847 

20. Clinopodes flavidus (C.L. Koch, 1847) 

Material examined. Malta: 1, Buskett, 30.12.1978, B/2, JS (MZ, MZ det.). 

General distribution. Poland, Czech Republic, Slovakia, Austria, Italy (incl. 
Sicily, excl. Sardinia), Slovenia, Croatia, Bosnia Herzegovina, Serbia, Montenegro, 
FYR Macedonia, Albania, Romania, Bulgaria, mainland and insular Greece (incl. 
Crete), Turkey, Cyprus, Palestine, Syria, European Russia, Crimea, Caucasus, 
Turkestan (Attems, 1929, 1949; Foddai et al., 1995; Kos, 1992; Orszâgh, 2001; Stoev, 
1997, 2001; Wytwer, 1997; Tajovsky, 2001; Zapparoli, 1995b, 1999). 

Chorotype. Turano-European (TUE). 

Ecological notes. The only locality from where this species was collected in the 
Maltese Islands is a semi-natural woodland. 

Tuoba Chamberlin, 1920 

2 1 . Tuoba poseidonis ( Verhoeff , 1 90 1 ) 

Material examined. Malta: 4, Golden Bay, 19.11.2001, AD (AM, AM det); 2, ibidem, 
19.11.2001, AD (AM. AM det). Gozo: 1. Xatt L-Ahmar, 1.5.2002, SS (AM. AM det); 1, Xatt 
L-Ahmar, 1.5.2002. ST (AM. AM det): 1. Xatt L-Ahmar, 1.5.2002, ST (AM. AM det); 25, Xatt 
L-Ahmar, 4.5.2002, AD (AM, AM det). 

General distribution. Coasts of S. France (incl. Corsica), Central and S. Italy 
(incl. Sardinia and Sicily), Slovenia, mainland and insular Greece, Jordan (Dead Sea), 



CENTIPEDES OF THE MALTESE ARCHIPELAGO 447 

Egypt (Red Sea), Somalia (Indian Ocean) (Attems, 1929; Brölemann, 1930; Foddai et 
al, 1995, 1996; Stoev, 1997; Zapparoli, 1990, 1991). 

Chorotype. Mediterranean (MED). 

Ecological notes. Found under stranded Posidonia debris on sandy beaches. 

FAUNISTIC AND ZOOGEOGRAPHIC REMARKS 

On the basis of the faunistic data available so far, twenty-one species of centi- 
pedes are here listed from the Maltese Archipelago, i.e. 1 Scutigeromorpha, 7 Litho- 
biomorpha, 3 Scolopendromorpha, and 10 Geophilomorpha. Of these, 20 have been 
confirmed to occur on the island of Malta, 1 1 species on Gozo and three on Cornino 
(Tab. I). 

The faunistic data presented in this study are significant, since they are the result 
of systematic collecting made in different seasons over a number of years and 
employing different collecting methods, but it is likely that the real number of species 
present in the Maltese Islands is slightly higher than reported here. 

One possible addition to the list may be Cryptops punicus Silvestri, 1896, a 
thermoxerophilous species widespread in the W-Mediterranean basin and known from 
the Tuscany Is. (Montecristo), Sardinia, Ustica, Sicily, Pantelleria, Lampedusa and 
Tunisia (cf. Zapparoli, 1995a). 

Other species that can be expected to occur in the Maltese Archipelago are 
Hydro schendyla submarina (Grube, 1872) and Nannophilus eximius (Meinert, 1870). 
H. submarina is a halophilous species, with a Mediterranean-Atlantic distribution for 
which some records are known for the Tyrrhenian (Tuscany Is., NE. Sardinia, 
Campania, Linosa: Zapparoli, 1995a) and Aegean areas (Zapparoli, 2002). N. eximius 
is a W-Mediterranean element, recorded from Macaronesia (Madeira, Canary Is.), N. 
Africa (Algeria, Tunisia) and S. Italy (Sicily, Calabria, Basilicata, Puglia); it is also 
present in the circum-Sicilian micro-insular systems (Egadi Is., Eolian Is., Pelagian 
Is.). 

Another species that may also be present is Geophilus insculptus Attems, 1895, 
a rather euryoecious geophilomorph widely distributed in Europe and also present in 
N-Africa. In the Tyrrhenian islands this species is known from many localities on 
Sardinia and Sicily and it has also been recorded from the Tuscany Is. (Giannutri, 
Giglio), Campane Is. (Ischia), Egadi Is., Eolian Is., Pantelleria and Lampedusa 
(Zapparoli, 1995a). 

With respect to our knowledge of the centipede fauna of individual Maltese 
islands, data are most complete for Malta, while the number of species that occur on 
Gozo and Cornino is very likely an underestimate. Some widespread Mediterranean 
elements (e.g.: Scutigera coleoptrata, Eupolybothrus nudicornis, Lithobius crassipes, 
Scolopendra oraniensis, Pachymerium ferrugineum, Tuoba poseidonis) have not been 
recorded from Gozo and Cornino, probably due to inadequate collecting on these 
islands. 

When compared to the faunas of the Central Mediterranean mainland and 
insular areas (Sicily, Eolian Is., Ustica, Egadi Is., Pantelleria, Linosa, Lampedusa and 
Tunisia), the centipede fauna of the Maltese Islands represents about a third of that 



448 M. ZAPPAROLI ET AL. 

known to date for this region as a whole (Tab. II). However, faunistic knowledge of 
these areas is still incomplete. Best known are the Sicilian (including the small 
surrounding islands) and Maltese faunas, both of which have been the object of recent 
and repeated studies. Just under fifty species have been recorded from these localities 
as a whole (Foddai et al., 1995, 1996). In contrast, knowledge of the centipedes of N. 
Africa, and particularly Tunisia, is much less complete and in need of updating. Some 
forty species have been recorded from this region to date, at least five of which are 
doubtful (Bròlemann, 1921, 1932; Zapparoli, unpublished records). 

Leaving out L. peregrinus, which is probably introduced, and L. forficatus, the 
presence of which on Malta is doubtful, almost 70% of the centipedes of the Maltese 
Islands are represented by species with a high dispersal ability. They are widespread in 
the Mediterranean area and are present in most of the aforementioned insular and main- 
land localities in the Central Mediterranean (Tabs II, III). 

Apart from Schendyla n. sp., the remaining Maltese centipede species are more 
or less widely distributed but absent from N. Africa (Lithobius lapidicola, L. trinacrius, 
Scolopendra oraniensis, Clinopodes flavidus) or from southern Italy {Bothriogaster 
signata). For Bothriogaster signata, however, the possibility that its presence on Malta 
is a result of anthropic introduction cannot be excluded. 

From a Zoogeographie point of view, an analysis of the chorotypes represented 
in the study area (Tab. Ill) shows the Maltese centipede fauna to have a strong 
"Mediterranean" character, with this chorological element approaching 67%. This 
agrees well with the results of studies on the centipede fauna of other insular systems 
of the Sicily Channel (Pantelleria, Linosa, Lampedusa) (Zapparoli, 1995a). Species 
with a wide distribution in the Holarctic Region and in Europe are poorly represented 
(22% and 11% respectively). 

Our studies on the centipede fauna of the Maltese Islands lead us to conclude 
that the composition of the fauna of these islands is affected mainly by ecological 
factors (the availability of suitable habitats) and by colonization-extinction events, 
rather than by paleogeographic and paleoclimatic factors. This is in accordance with 
the conclusions of previous studies on the centipedes of the W-Mediterranean micro- 
insular systems (Foddai et al., 1996). 

ACKNOWLEDGEMENTS 

We wish to thank the many people who over the years collected or donated the material 
on which this paper is based. We are also indebted to Volker Mahnert and Peter 
Schwendinger, Director and Curator of the Muséum d'histoire naturelle, Genève, for a 
careful reading of the manuscript. PJS was supported by research grants awarded by 
the University of Malta Research Committee, for which he is grateful. 

REFERENCES 

Alexander, D. 1988. A review of the physical geography of Malta and its significance for 

tectonic geomorphology. Quaternary Science Reviews 7: 41-53. 
Attems, CG. 1907. Myriopoden aus der Krim und dem Kaukasus von Dr. A. Stuxberg 

gesammelt. Archiv för Zoologi 25: 1-16 + 2 pis. 



CENTIPEDES OF THE MALTESE ARCHIPELAGO 449 



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Tab. I - Distribution of the centipedes recorded from the main islands of the Maltese 
Archipelago. Symbols: + = present; - = absent; ? = doubtful record; * = introduced or probably 
introduced species. Only confirmed species are considered in the total. 

Malta Gozo Cornino 

Scutigera coleoptrata (Linné, 1758) + - - 

Eupolybothrus (Allopolybothrus) nudicornis (Gervais, 1837) + + - 

Lithobius (Lithobius) castaneus Newport, 1 844 + - - 

Lithobius (Lithobius) forficatus (Linné, 1758) ? - - 

Lithobius (Lithobius) lapidicola Meinert, 1 872 + - - 

Lithobius (Lithobius) peregrinus Latzel, 1880 * 

Lithobius (Lithobius) trinacrius Verhoeff, 1925 + + - 

Lithobius (Monotarsobius) crassipes L. Koch, 1 862 + + 

Scolopendra cingulata Latreille, 1829 + + + 

Scolopendra oraniensis Lucas, 1846 + + - 

Cryptops trisulcatus Bròlemann, 1902 + + + 

Himantarium gabrielis (Linné, 1767) + - - 

Stigmatogaster gracilis (Meinert, 1870) + - - 

Bothriogaster signata (Kessler, 1874) * * * 

Dignathodon microcephalus (Lucas, 1846) + + - 

Henia (Meinertia) bicarinata (Meinert, 1870) + + - 

Henia (Chaetechelyne) vesuviana (Newport, 1845) + - - 

Schendyla sp. n. + - • - 

Pachymerium ferrugineum (C.L. Koch, 1835) + + - 

Clinopodes flavidus C.L. Koch, 1847 + - - 

Tuoba poseidonis (Verhoeff, 1901) + + - 

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2 



REVUE SUISSE DE ZOOLOGIE 
Tome 1 1 1 — Fascicule 2 

Pages 

Schätti, Beat & Monsch, Paul. Systematics and phylogenetic relationships 
of Whip snakes {Hierophis Fitzinger) and Zamenis andreana Werner, 
1917 (Reptilia: Squamata: Colubrinae) 239-256 

Hou, Zhong-e, Li, Shuqiang & Platvoet, Dirk. Three new species of the 
genus Gammarus from tributaries of the Hi River, China (Crustacea, 
Amphipoda, Gammaridae) 257-284 

Puthz, Volker. Neue orientalische Arten der Gattung Stenus Latreille aus 
dem Genfer Museum (Coleoptera: Staphylinidae). 284. Beitrag zur 
Kenntnis der Steninen 285-301 

Benjamin, Suresh P. Nesticeila marapu sp. n., a blind nesticid (Araneae: 

Nesticidae) from Sumba, Indonesia 303-307 

Bollache, Loïc. Dìkerogammarus villosus (Crustacea: Amphipoda): 

another invasive species in Lake Geneva 309-313 

Schuchert, Peter. Revision of the European athecate hydroids and their 
medusae (Hydrozoa, Cnidaria): Families Oceanidae and Pachycor- 
dylidae 315-369 

De Chambrter, Alain. Redescription of Ophiotaenia hylae Johnston, 1912 
(Eucestoda: Proteocephalidea), parasite of Litoria aurea (Amphibia: 
Hylidae) from Australia 371-380 

Pœrallini, Riccardo, Keller, Albert & Moretti, Marco. Chiave di deter- 
minazione dei Chirotteri (Mammalia) della Svizzera attraverso l'osser- 
vazione al microscopio ottico della struttura dei peli 381-393 

Baur, Hannes, Landau Lüscher, Isabelle, Müller Gabi, Schmidt, 
Marcus & Coray, Armin. Taxonomie der Bernstein-Waldschabe Ecto- 
bius vittiventris (A. Costa, 1847) (Blattodea: Blattellidae) und ihre 
Verbreitung in der Schweiz 395-424 

Ng, Heok Hee & Kottelat, Maurice. Amblyrhynchichthys micracanthus, a 
new species of cyprinid fish from Indochina (Cypriniformes: Cypri- 
nidae) 425-432 

Zapparoli, Marzio, Minelli, Alessandro & Schembri, Patrick J. The 

centipedes of the Maltese Archipelago (Chilopoda) 433-456 



REVUE SUISSE DE ZOOLOGIE 
Volume 1 1 1 — Number 2 

Pages 

Schätti, Beat & Monsch, Paul. Systematics and phylogenetic relationships 
of Whip snakes (Hierophis Fitzinger) and Zamenis andreana Werner, 
1917 (Reptilia: Squamata: Colubrinae) 239-256 

Hou, Zhong-e, Li, Shuqiang & Rlatvoet, Dirk. Three new species of the 
genus Gammarus from tributaries of the Hi River, China (Crustacea, 
Amphipoda, Gammaridae) 257-284 

Puthz, Volker. New Oriental species of the genus Stenus Latreille from the 
Geneva Museum (Coleoptera: Staphylinidae). 284 th Contribution to 
the knowledge of Steninae 285-301 

Benjamin, Suresh P. Nesticella marapu sp. n., a blind nesticid (Araneae: 

Nesticidae) from Sumba, Indonesia 303-307 

Bollache, Loïc. Dikerogammarus villosus (Crustacea: Amphipoda): 

another invasive species in Lake Geneva 309-313 

Schuchert, Peter. Revision of the European athecate hydroids and their 
medusae (Hydrozoa, Cnidaria): Families Oceanidae and Pachycor- 
dylidae 315-369 

De Chambrier, Alain. Redescription of Ophiotaenia hylae Johnston, 1912 
(Eucestoda: Proteocephalidea), parasite of Litoria aurea (Amphibia: 
Hylidae) from Australia 371-380 

Pierallini, Riccardo, Keller, Albert & Moretti, Marco. Identification key 
of bats (Chiroptera) from Switzerland based on hair structure observed 
through optical microscope 381-393 

Baur, Hannes, Landau Lüscher, Isabelle, Müller Gabi, Schmidt, 
Marcus & Coray, Armin. Taxonomy of the field-dwelling cockroach 
Ectobius vittiventris (A. Costa, 1847) (Blattodea: Blattellidae) and its 
distribution in Switzerland 395-424 

Ng, Heok Hee & Kottelat, Maurice. Amblyrhynchichthys mìcracanthus, a 
new species of cyprinid fish from Indochina (Cyprinif ormes: Cypri- 
nidae) 425-432 

Zapparoli, Marzio, Minelli, Alessandro & Schembri, Patrick J. The 

centipedes of the Maltese Archipelago (Chilopoda) 433-456 



Indexed in Current Contents, Science Citation Index 



PUBLICATIONS DU MUSEUM D'HISTOIRE NATURELLE DE GENÈVE 

CATALOGUE DES INVERTÉBRÉS DE LA SUISSE, N«s 1-17 (1908-1926) série Fr. 285.- 

(prix des fascicules sur demande) 

REVUE DE PALÉOBIOLOGIE Echange ou par fascicule Fr. 35.— 

LE RHINOLOPHE (Bulletin du centre d'étude des chauves-souris) par fascicule Fr. 35.— 

THE EUROPEAN PROTURA: THEIR TAXONOMY, ECOLOGY AND 
DISTRIBUTION, WITH KEYS FOR DETERMINATION 
J. Nosek, 345 p., 1973 Fr. 30.— 

CLASSIFICATION OF THE DIPLOPODA 

R. L. Hoffman, 237 p., 1979 Fr. 30.— 

LES OISEAUX NICHEURS DU CANTON DE GENÈVE 
P. Géroudet, C. Guex & M. Maire 
351 p., nombreuses cartes et figures, 1983 Fr. 45. — 

CATALOGUE COMMENTÉ DES TYPES DECHINODERMES ACTUELS 
CONSERVÉS DANS LES COLLECTIONS NATIONALES SUISSES, 
SUIVI D'UNE NOTICE SUR LA CONTRIBUTION DE LOUIS AGASSIZ 
À LA CONNAISSANCE DES ECHINODERMES ACTUELS 
M. Jangoux, 67 p., 1985 Fr. 15.— 

RADULAS DE GASTÉROPODES LITTORAUX DE LA MANCHE 
(COTENTIN-BAIE DE SEINE, FRANCE) 
Y. Finet, J. Wüest & K. Mareda, 62 p., 1991 Fr. 10.— 

GASTROPODS OF THE CHANNEL AND ATLANTIC OCEAN: 
SHELLS AND RADULAS 
Y. Finet, J. Wüest & K. Mareda, 1992 Fr. 30. — 

O. SCHMIDT SPONGE CATALOGUE 

R. Desqueyroux-Faundez & S.M. Stone, 190 p., 1992 Fr. 40.— 

ATLAS DE RÉPARTITION DES AMPHIBIENS 
ET REPTILES DU CANTON DE GENÈVE 
A. Keller, V. Aellen & V. Mahnert, 48 p., 1993 Fr. 15. — 

THE MARINE MOLLUSKS OF THE GALAPAGOS ISLANDS: 
A DOCUMENTED FAUNAL LIST 
Y. Finet, 180 p.. 1995 Fr. 30.— 

NOTICE SUR LES COLLECTIONS MALACOLOGIQUES 
DU MUSEUM D'HISTOIRE NATURELLE DE GENEVE 
J.-C. Cailliez, 49 p., 1995 Fr. 22.— 

PROCEEDINGS OF THE XHIth INTERNATIONAL CONGRESS 

OF ARACHNOLOGY, Geneva 1995 (ed. V. Mahnert), 720 p. (2 vol.), 1996 Fr. 160.— 

CATALOGUE OF THE SCAPHIDIINAE (COLEOPTERA: STAPHYLINIDAE) 

(Instrumenta Biodiversitatis I), I. LObl, xii + 190 p., 1997 Fr. 50. — 

CATALOGUE SYNONYMIQUE ET GÉOGRAPHIQUE DES SYRPHIDAE (DIPTERA) 
DE LA RÉGION AFROTROPICALE 
(Instrumenta Biodiversitatis II), H. G. DlRlCKX, x +187 p., 1998 Fr. 50.— 

A REVISION OF THE CORYLOPHIDAE (COLEOPTERA) OF THE 
WEST PALAEARCTIC REGION 
(Instrumenta Biodiversitatis III), S. Bowestead, 203 p., 1999 Fr. 60.— 

THE HERPETOFAUNA OF SOUTHERN YEMEN AND THE 
SOKOTRA ARCHIPELAGO 

(Instrumenta Biodiversitatis IV), B. SCHÄTTI & A. Desvoignes, 
178 p., 1999 Fr. 70.— 

PSOCOPTERA (INSECTA): WORLD CATALOGUE AND BIBLIOGRAPHY 

(Instrumenta Biodiversitatis V), C. Lienhard & C. N. Smithers, 

xli + 745 p., 2002 Fr. 180.— 

REVISION DER PALÄARKTISCHEN ARTEN DER GATTUNG BRACHYGLUTA 
THOMSON, 1859 (COLEOPTERA, STAPHYLINIDAE) (1. Teil) 
(Instrumenta Biodiversitatis VI), G. Sabella, Ch. Buckle, V. Brachat 
& C. Besuchet, vi + 283 p., 2004 Fr. 100.— 



Volume 1 1 1 - Number 2 - 2004 

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Penard, E. 1888. Recherches sur le Ceratium macroceros. Thèse, Genève, 43 pp. 
Penard, E. 1889. Etudes sur quelques Héliozoaires d'eau douce. Archives de Biologie 9: 1-61. 
Mertens. R. & Wermuth, H. 1960. Die Amphibien und Reptilien Europas. Kramer. Frankfurt am Main. XI + 264 pp. 
Handley, C. O. Jr 1966. Checklist of the mammals of Panama (pp. 753-795). In: Wenzel, R. L. & Tipton. V. J. 

(eds). Ectoparasites of Panama. Field Museum of Natural Histoiy, Chicago, XII + 861 pp. 

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